Palaeopopulations of Late Pleistocene Top Predators in Europe: Ice Age Spotted Hyenas and Steppe Lions in Battle and Competition about Prey

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Palaeopopulations of Late Pleistocene Top Predators in Europe: Ice Age Spotted Hyenas and Steppe Lions in Battle and Competition about Prey

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Palaeopopulations of Late Pleistocene Top Predators in Europe: Ice Age Spotted Hyenas and Steppe Lions in Battle and Competition about Prey
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Cajus G. DiedrichPrivate Research Institute Paleologic, Petra Bezruce 96, 26751 Zdice, Czech Republic Correspondence should be addressed to Cajus G. Diedrich; cdiedri@gmx.net Received 21 March 2013; Accepted 11 June 2013 Academic Editor: Vlad Codrea Copyright (c) 2014 Cajus G. Diedrich. This is an open access article distributed under the Creative Commons Attribution License, which permits unrestricted use, distribution, and reproduction in any medium, provided the original work is properly cited. Late Pleistocene spotted hyena Crocuta crocuta spelaea (Goldfuss, 1823) and steppe lion Panthera leo spelaea (Goldfuss, 1810) were top predators in Central Europe. The fossil record (2.303 hyena/1.373 lion bones = ratio 3/1) from 106 cave and open air sites demonstrates comparable associations to modern African hyenas/lions resulting in competition about prey and territory. Cannibalism within extinct spotted hyenas is well documented, including two individual skeletons. Those hyenas produced bone accumulations at dens. Feeding specializations on different megamammal groups are demonstrated for Late Pleistocene hyenas whose prey partly overlaps (e.g., cave bears) with those of lions and wolves. At most fossil sites, 1-3% of the lion remains indicate scavenging of lions by hyenas. The larger Late Pleistocene felids focussed on cervids (reindeers specialization during the high glacial = LGM), on bovids (steppe bison/aurochs), and possibly on saiga antelope and on the cave bear, hunting deep in caves during their hibernations and targeting cubs. The cave bear feeding was the target of all three top predators (lions, hyenas, and wolves) in the Late Pleistocene boreal forests which caused deathly conflicts in caves between them, especially with lions/hyenas and herbivorous cave bears that have no modern analogue.
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Cajus G. DiedrichPrivate Research Institute
Paleologic, Petra Bezruce 96, 26751 Zdice, Czech Republic
Correspondence should be addressed to Cajus G. Diedrich;
cdiedri@gmx.net Received 21 March 2013; Accepted 11 June 2013
Academic Editor: Vlad Codrea Copyright 2014 Cajus G.
Diedrich. This is an open access article distributed under
the Creative Commons Attribution License, which permits
unrestricted use, distribution, and reproduction in any
medium, provided the original work is properly cited. Late
Pleistocene spotted hyena Crocuta crocuta spelaea (Goldfuss,
1823) and steppe lion Panthera leo spelaea (Goldfuss, 1810)
were top predators in Central Europe. The fossil record
(2.303 hyena/1.373 lion bones = ratio 3/1) from 106 cave and
open air sites demonstrates comparable associations to modern
African hyenas/lions resulting in competition about prey and
territory. Cannibalism within extinct spotted hyenas is well
documented, including two individual skeletons. Those hyenas
produced bone accumulations at dens. Feeding specializations
on different megamammal groups are demonstrated for Late
Pleistocene hyenas whose prey partly overlaps (e.g., cave
bears) with those of lions and wolves. At most fossil sites,
1-3% of the lion remains indicate scavenging of lions by
hyenas. The larger Late Pleistocene felids focussed on
cervids (reindeers specialization during the high glacial =
LGM), on bovids (steppe bison/aurochs), and possibly on saiga
antelope and on the cave bear, hunting deep in caves during
their hibernations and targeting cubs. The cave bear feeding
was the target of all three top predators (lions, hyenas, and
wolves) in the Late Pleistocene boreal forests which caused
deathly conflicts in caves between them, especially with
lions/hyenas and herbivorous cave bears that have no modern
analogue.



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HindawiPublishingCorporation PaleontologyJournal Volume!"#$,ArticleID#"%!"&, &$ pages http://dx.doi.org/#".##''/!"#$/#"%!"& ResearchArticle PalaeopopulationsofLatePleistoceneTopPredatorsinEurope: IceAgeSpottedHyenasandSteppeLionsinBattleand CompetitionaboutPrey CajusG.Diedrich PrivateResearchInstitutePaleologic,PetraBezruce!",#"$%&Zdice,CzechRepublic CorrespondenceshouldbeaddressedtoCajusG.Diedrich;cdiedri@gmx.net Received!#March!"#&;Accepted##June!"#& AcademicEditor:VladCodrea Copyright!"#$CajusG.Diedrich.(isisanopenaccessarticledistributedundertheCreativeCommonsAttributionLicense, whichpermitsunrestricteduse,distribution,andreproductio ninanymedium,providedtheoriginalworkisproperlycited. LatePleistocenespottedhyena Crocutacrocutaspelaea (Goldfuss,#)!&)andsteppelion Pantheraleospelaea (Goldfuss,#)#") weretoppredatorsinCentralEurope.(efossilrecord(!.&"&hyena/#.&*&lionbones=ratio&/#)from#"%caveandopenair sitesdemonstratescomparableassociationstomodernAfricanhyenas/lionsresultingincompetitionaboutpreyandterritory. Cannibalismwithinextinctspottedhyenasiswelldocumented,includingtwoindividualskeletons.(osehyenasproducedbone accumulationsatdens.Feedingspecializationsondi+erentmegamammalgroupsaredemonstratedforLatePleistocenehyenas whosepreypartlyoverlaps(e.g.,cavebears)withthoseoflionsandwolves.Atmostfossilsites,#&%ofthelionremainsindicate scavengingoflionsbyhyenas.(elargerLatePleistocenefelidsfocussedoncervids(reindeersspecializationduringthehighglacial =LGM),onbovids(steppebison/aurochs),andpossiblyonsaigaantelopeandonthecavebear,huntingdeepincavesduringtheir hibernationsandtargetingcubs.(ecavebearfeedingwasthetargetofallthreetoppredators(lions,hyenas,andwolves)inthe LatePleistoceneborealforestswhichcauseddeathlycon,ictsincavesbetweenthem,especiallywithlions/hyenasandherbivorous cavebearsthathavenomodernanalogue. 1.Introduction PleistocenetoppredatorresearchstartedinEuropewiththe rstdiscoveriesof"foreignanimalnds"intheZoolithen Cave(Germany)in#**$whentheGermanPriestEsper discoveredsomehyena(Figures # & ),lion(Figures $ % ), wolf,andcavebearremainsandexplainedthemresulting fromthe"greatdeluge"[ # ].Rosenm ullercollectedalarge amountofcavebearremainsincludingthe Ursusspelaeus holotypeskull[ ]andthelargestamountoflionmaterialin thesamecave[ & ].( iscollectionsurviveduntiltoday[ ]. ( eFrenchzoologistCuvierinterpretedin#)"'[ $ ]some skullfragmentsfromtheGermanZoolithenCaveas"hyena". ItwastheGermanPaleontologistGoldfusswhodescribed andnamedthe-rsttoppredatorofthePleistocenewiththe holotypeskullofthe"cavelion Felisspelaea "in#)#"[ % ]. In#)!&hepublishedonthesecondlargepredatorwiththe holotypeskullofthe"cavehyena Hyaenaspelaea "[ ].Both oftheseskullsfromtheZoolithenCaveweredescribedin moredetaila.ertheirrediscoveryin!""/[ ) ].Finally,also the"cavewolf Canisspelaeus "wasdescribedin#)!&basedon acubskullfoundalsointheZoolithenCave[ ]. ( emoderneraof"cavehyena"denresearchstartedin #)!&,whentheEnglishGeologistBucklandpublishedhis "ReliquiaeDiluvianae"[ / ].Modernhyenadeninterpretationfromhistorictimeswasbasedonmainlythefamous Kent'sCaverns(E)andtheK onig-LudwigsCave(D,there mentionedas"KuhstallorRabensteinCave")[ / ].( eGermanBiologistGiebelexcavatedhyena[ #" ]andotherfaunal remainsfromPerickCaves(=SundwigCave,Sauerland Karst)in#)$/#)'!inGermanyandhyenaandpreyfauna remainsfromtheSewecken-BergeandWesteregeln(D)[ ## ] openairgypsumkarstareasofnorthern-centralGermany.In #)%&,Dawkinsdiscoveredthehyenabone-rich($%*remains, mainlyteeth)andNeanderthalartefactbearing(e.g.,bifaces) WookeyHoleCavehyenadenandoverlappingPleistocene Neanderthalhumancampsite(E)[ #! #& ].Denresearch continuedwiththedescriptionsofthefaunasoffamoushyena

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! PaleontologyJournal V!pustek Cave (CZ) Koneprusy Cave (CZ) Srbsko Chlum-Kom’n Cave (CZ) 500 km London Berlin Paris Warsaw Oslo Bukarest Rome Athene Barcelona Madrid Munich Amsterdam Budapest Glacier Hyena absencesmall cave bears Glaciers Crocuta crocuta spelaea Tundra Forest tundra Boreal conifere forests Temperate broadleaf forests Mixed forests Taiga steppe Mediterranean forests Mountains with few vegetation Skeletons, skeleton remains Zoolithen Cave (D) Perick Caves (D) Westeregeln (D) Lindenthal Cave (D) Sloup Cave (CZ) V!pustek Cave (CZ) Koneprusy Cave (CZ) Srbsko Cave (CZ) Oase Cave (Ro) 500 km Carpathians Dinarids London Berlin Paris Warzaw Oslo Moskow Bukarest Rome Athene Barcelona Madrid Munich Budapest Glacier Sandfort Hill Cave (GB)Zoolithen Cave (D) Westeregeln gypsum karst (D) Sloup Cave (CZ) V!pustek Cave (CZ) Srbsko Cave (CZ) Skull shape types Flat Sligth convex Strong convex Oase Cave (Ro) Bottrop Open air (D) Amsterdam iede gypsum karst (D) Pšssneck Cavity (D) Hohenmšlsen open air (D) Kšnigsaue open air (D) Pin Hole Cave (GB) Bad Wildungen open air (D) BrŸhl-Spielewiesen open air (D) Ketsch-Kreuzwiese open air (D) Erkenbrechtsweiler limestone karst (D) Irpfel Cave (D) Main study area Perick Caves (D) Trmice open air (CZ) Javorka Cave (CZ) Teufelskammer Cave (D) Badel Cave (A) (b)(c) (d) Hutton Cave (GB) Wookey Hole Cave (GB) San Teodoro Cave (It) Bleadon Cave (GB) Townton Cave (GB) Agios Georgios Cave (Gr) Sophie's Cave (D) Plumettes Cave (F) Trou du Cluzeau Cave (F) Rochelot Cave (F) Sandfort Hill Cave (GB) Pin Hole Cave (GB) Crocuta crocuta crocuta Late Pleistocene (113,000-24,000 BP) (a) Crocuta crocuta spelaea Alps Carpathians Dinarids Alps F01234#:(a)SpottedhyenasgloballyduringtheLatePleistocene.((b)and(c))IceAgespottedhyena Crocutacrocutaspelaea (Goldfuss, #)!&)sitesinEuropeduringtheLatePleistocenebeforetheLastGlacialMaximum(##&."""!$."""BP)withabsenceinalpineregions(= redareas).(d)SkeletonsfromEurope:adultindividualskeletonfromV ypustekCave(MoravianKarst,CZ,coll.NHMW);composedcub skeletonfromSrbskoChlum-Kom nCave(BohemianKarst,coll.NHMP);adultindividualskeletonfromKon eprusyCave(BohemianKarst, CZ,coll.NHMP).

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PaleontologyJournal & dencavesitessuchastheLindentalCaveGera(D)in#)%*, wherephosphaticexcrementlayerswerereportedandwhere fracturedrhinocerosbonesresultingfromhyenashavebeen misidentiedas"PleistoceneNeanderthaltools"[ #$ ].( is smallcavealsohadartefactsfromNeanderthals,whichlead tohistoricalconclusionsthatthesenontoolbonesmustbe the"kitchenrubbish"ofPleistocenehumanswho"chewedon thebones"leavingbitemarks(="Osteodontokeraticculture" [ #' ]).NehringcontinuedtheGermanLatePleistocenehyena researchandcollectedsomeskullsandseveralpostcranial hyenaremainsandpreybonesfromanewopenairgypsum karstdensite(iede(D)[ #% ]andattheR osenbeckCave(D) [ #* ].In#)/!Haarl estartedthe"hyenaden"researchinSEFrance[ #) ].Reynoldspublishedin#/"![ #/ ]byfarthebest monographontheEnglishLatePleistocenehyenaremains ofitstime,butheconfusedsomematerialas"individual skeletons"whichareactuallyfromatleasttwodi+erentcaves, WookeyHole[ #/ !" ]andSandfortHill[ #/ !" ]Caves(E). Tworediscoveredskulls[ #* ]fromthesetwolocalitiesanda largecollectionwhichwashiddenduringthesecondWorld WarundercoalsintheSomersetMuseum[ !" ]havenot beenanalysedyetatpresent.(eTeufelsluckenCave(A) wasanotherhyenacavedenandtherstdescribedfrom Austriain#/&*[ !# ].Zapfepresentedin#/&/apaperon damagedbonesbyhyenasfromdi+erentcaveandopenair sites,especiallyfromAustria[ !! ].A.ertheSecondWorld War,newLatePleistocenehyenadenswerediscoveredin#/'* withtheTorbryanCaveandTonewtonCave(E)[ !" !& ].In #/%#,(eniuspresentedhyenadamagedandchewedbones fromAustriaandcriticallydiscussedthe"Osteodontokeratic culture"oftheTertiaryAustralopithecines(Hominidae), whichwerethoughtincorrectlytohaveproducedhominid bitedamageonbones,whichwereindeedmadebyhyenas [ #' ].(eCzechMoravianPaleontologistMusilpresenteda furtherimportantmonographaboutthehyenasandtheir preyfromtheSveduvSt ulCavehyenaden(MoravianKarst, CZ)in#/%![ !$ ].In#/%)somearticulatedhyenaprey remainsfromopenairloesssites(=mammothsteppe)were mentionedfromthesiteofAchenheim(D)[ !' ],whichconvincinglydemonstratedhyenaactivitiesoutsidethecavesfor the-rsttime.AnothersouthernGermanhyenadencave,the Au5auserCave,was#/)!shortlypresentedbutremainednot analysed[ !% ].In#/)&,thersthyenaremainswererecovered withotherLatePleistocenebonesonthesea, ooroftheNorth Seaby-shingboats[ !* ].In#/)/,theLatePleistocenehyena denCamiacCave(F)waspresented[ !) ]andin#//!another importanthyenadenfromAgiosGeorgiosCave(Gr)[ !/ ]. ( elattercavecontainedmanyLatePleistocenehorseand donkeyremainsthatwerethemainpreyofhyenasinacub raisingdensite,butthiswasnotwellanalysedin"hyena dencontext."In#//%inSW-France,especiallyTournepiche beganstudieson"boneassemblages"toseparatetheirhuman fromcarnivoregenesis(oroverlap)mainlyatcavesites.He identied#%LatePleistocenehyenacavedensbutdidnot followqualitativepreyorpopulationstructureanalyses[ &" && ].( eseincludethecoldperiodhyenapreyfaunafrom TrouduCluzeauCave[ &" ],PlumettesCave[ &" ],andthe EemianwarmperiodpreyfaunafromRochelotCave[ &$ ]. Inthelatterhyenaden,evenNeanderthalhumanremains (teethandcrushedlongbonefragments)werefoundwithin thebovid/equid/suiddominatedpreyboneaccumulation [ &$ ].(emosteasternknownLatePleistocenehyenaden wasreportedin!"""fromtheProlomIICave[ &' ]in theCrimea,wheretherearehyenadensoverlappingwith humancampsitesatcaveentrances/portals.(eoverlapping o.endoesnotenableclearattributiontohyenapreyor humankitchenrubbish.Interestingly,otherhyenadenswith possiblyoverlappingwolfdenswerepublishedforsomeLate PleistoceneItaliancavesin!""$[ &% ].Mostprobablyinstead thewolveswerealsopreyandimportedbyhyenas,which isunsolvedyet.( eSanTeodoroCaveonSicily(I)was thenpresentedin!"##,demonstratingthelowmarinesea levelduringtheglacialoftheLatePleistocenewhichallowed themigrationofhyenastotheislandofpresentday[ &* ]. Currantcompiledseveralforgottenandnewhyenadens fromEnglandin!""$,includingtherediscoveredReynolds collectionsfromWorldWarIIdescribingfurthercavesas hyenadenssuchasBleadon,Cavern,orHuttonCavern [ !" ].AskullfromPinHole[ #* ]wasaddedforEnglish sites.In!"")Polishhyenaremainswerecompiled[ &) ].In theCzechRepublicin!""$,theauthor's"EuropeanLate Pleistocenespottedhyenaproject"providedanoverviewof LatePleistocenehyenadentypes,mainlycaves,whichwas presentedfortheBohemianKarstmountainousregion(= between#'"and%'"a.s.l.)nearPrague[ &/ ].Inaddition themostfamousCzechLatePleistocenecavesandhyena remains,and-rstindividualhyenaskeletons,werereported fromtheKon eprusyCave[ $" ]andSrbskoChlum-Kom n Cave[ $# ].Recenthyenapopulationsandpreyanalysesbased onmodernAfricanspottedhyenaethologywereusedfor thefamousSloupCave[ $! ]andtheV ypustekCave[ $! ] intheMoravianKarst(CZ).InGermany,severalimportant forgottenoroverlookedhyenadencave(#"sites)andopenair sites(!sites)havebeendescribedinthepastdecade.(ese includetheSauerlandKarstcave-richregioninnorthwestern Germanyand*'"bonesand#&skullsmainlyfromthePerick Caves[ #" ],TeufelskammerCave[ $& ],BalveCave[ $$ ],and R osenbeckCave[ #* ].( emostimportantZoolithenCave intheGermanFranconianKarstwasalsoreviewedforits hyena[ $' ]andhyenapreyandlion[ $% ]content,whereas theFranconianSophie'sCave[ $* ]inthatregionsupported thecomplextaphonomysolvingbetweentoppredatorsand theirscavengingandhuntofcavebearsinEurope.To understandthelifeofextinctLatePleistocenespottedhyenas outsidethecavesinlowlandsoftheUpperRhineValley [ $) ],isolatedremainswerepresentedfromriverterrace gravelpitsitessimilartotheM unsterlandBaylowlands[ $/ ], whereasapopulationanditspreyfromtheriverterracesite Bottrop[ '" ]isakeysitewithmorethen&."""bonesto understandopenairboneaccumulationsordensandlifeof LatePleistocenespottedhyenas,especiallyalongrivers(river terracedentypes).(ere,bonesfromgravelpitsiteshave beensimplyincorrectlyattributedto",uvialtransport,"but perfectpreservedbitedamageandhighamountsofchewed largemammalbones,suchasthosefoundinBottroporSelmTernscheandHernealongtheLippeandEmscherRivers, demonstratethattheseboneaccumulationsaresimilaras thoseinmodernAfricanfromspottedhyenasalongwater

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$ PaleontologyJournal bodies.OtherGermanopenairhyenadenandboneaccumulationsitesalongancientriversandinloessmammothsteppe palaeoenvironmentshavebeenpublished.(eseincludeBad Wildungen(therehyenabirthden=recycledbadger/foxdens inloess)[ '# ]andK onigsaue(overlappingNeanderthalcamp site)[ '! ],whereasmorerecentstudieshaveanalyzedopenair gypsumkarst(Westeregeln[ '& ],Sewecken-Berge[ '$ ])and limestonekarstdens(FuchslukenCavity[ '' ],W usteScheuer Cavity[ '% ])tocoverallmorphologicaldentypes,especially thoseoutsidecaves.Hyenascavengingsiteswithremains oftheirlargestprey(woollyrhinoceroscarcass-Kr olpa[ '% ], woollymammothcarcass-Siegsdorf[ '* ],andforestelephant carcasses-Neumark-NordLake#[ ') ])demonstratethetop predatorbutcheringtechniqueanddeathlycon, ictwithlions overprey. LatePleistocenelionsweredescribedforthe-rsttime in#)#"basedonaskull( Figure' )fromtheZoolithenCave hyenaandcavebearden(D)[ ].Becausetheseinitialnds werefromcaves,theywerehistoricallynamed"cavelions," whichwasrevisedrecentlyto"steppelion"[ ) ].In#/"",the mostnorthernsingletoothandboneremainsoflionsfrom Englishhyenadencaveswerepresentedinamonograph[ %/ ]. In#/"%,"cavelion"remainsfromFrancewerepublished[ *" ]. AsingleskullwasthenreportedfromtheGentherCave,a cavebeardeninUpperFranconia(D)in#/'&[ *# ].In#/%),a discussionbetweenthecomparisonoflionremainsfromthe German/European"Pleistocene"andmodernAfricanlions resultedinthedeterminationthatthe"Pleistocene"forms(it wasgeneralized)musthavebeenslightlylarger(#/$)than modernones[ *! ],whichisthetruthonlyfortheMiddle Pleistocene(Saalianglacial) P.leofossilis subspecies,indeed. In#/'*,LatePleistocenelionswerethenattributedtothe modernlionsubspecies,as P.leospelaea ,osteometrically basedonnewmaterialfromFrance[ *& ].In#/)&,-shingboats recoveredmanyPleistocenemammalbones,includinglion remains,onthesea, ooroftheNorthSea[ !* ].( erstLate Pleistoceneindividuallionskeletonwasthendiscoveredin Arrikrutz(Es)in#/)#[ *$ ],withanadditionalskeletoninAustria[ *' ].Lionskullsandpostcranialremainswerepublished byArgantin#/))fromthecavebeardenAzeCave(F),which containedMiddle/LatePleistocenematerialmixed,including someMiddlePleistocene P.leofossilis remains[ *% ].Another skeletonfromalargemalelionwasfoundbefore#//!beside amammothcarcassattheopenairsiteofSiegsdorf(D) [ ** ].Cranialandpostcranialbonesfollowedfromacave excavationintheGermancavebeardensiteofHermann's Cave[ *) ].FromCroatia,lionswerecompiledwithsingle remainsfromseveralcavesiteswithoutcleartaphonomic context[ */ ].FurthertotheeastinYakutia,the-rst"cave lions"(noteven"steppelions,"insteadanothersubspecies P.leovereshchagini )werereportedfromopenairlocalities [ )" ].Anothersingle P.l.spelaea lionskullwasrecovered inZandobbio(It)datingtotheEemianinterglacial[ )# ]. FromCzechRepublic,remainswerecompiledin!""*from severalopenair(e.g.,Praha)andhyenadencavesites[ %& ]. In!""),anoverviewofPolishlionremainswaspresented, againwithoutaclearunderstandingoftheirtaphonomy[ &) ]. ( erstEuropean"skeletons"werepublishedfromtheSloup Cavehyenaandcavebearden(CZ),whichnewstudies revealedbothcompositeskeletonsofdi+erentindividuals [ %! ].(eonlyindividualskeletonisfromtheSrbskoChlumKom nCave,representinganearlyadultlioness( Figure$ ) withevidenceofabraincasetraumainjuredanimal[ %! ]. ( isskeletonisimportantforunderstandingthecon, ict betweenbothtoppredators,especiallyatcavedensites. Severalskullsandpostcranialremainswereanalyzedfor sexualdimorphismandtaphonomiccontext,especiallyfrom cavebearandhyenadensitesinGermany,whereforthe -rsttimethehyena-lionantagonismandscavengingof lioncarcassesbyhyenaswerediscussed[ %$ )! )& ].( e mostimportantnewdiscoveriesin!""/werethreelion skeletonsindi+erenttaphonomicsituationsfounddeeply withintheentranceofacavebearhibernationplateausin theRomanianUrs ilorCave[ )$ ].( isdiscoveryyieldeda clearevidencefortheactivehuntingofhibernatingcave bearsbylionsdeepinlargerandcomplexcavesystems[ %' ]. Furtherimportantistheopenairsitediscoveryofadiseased olderlioness( Figure$ )inbetweentheEemianinterglacial elephantgraveyardNeumark-NordLake#.(isskeletonwas interpretedtohaveresultedfromahyena/lioncon, ictover theirlargestprey[ %# ].(eincorporationoflionremainsat overlappinghyenadenandhumancampsiteswasdiscussed criticallyfortheBalveCave(D)[ )' ],whereitwasincorrectly believedthatNeanderthals/AurignacianmodernPleistocene LatePalaeolithichumansimported"lionfurs"a.ertheir killstothecampsite.Indeed,thoselionremainswerelater demonstratedtohaveresultedfromhyenaandcavebear con, icts.Alistingandanalysisofsomematerialwithoutclear taphonomy,includingasingleskull(Be se nov aCave),were compiledfromtenSlovakiancaves[ )% ].( elargestEuropean LatePleistocenelionfossilrecord(=palaeopopulation)was recentlydescribedin!"##forZoolithenCave,whereevidence forcavebearhuntingbylionswasestablishedbytaphonomic studies[ $$ ].Openairlionmaterialwasthenreviewedfor northernGermanyforallkindofsites( Figure$ ),ofwhich severalboneshadalsobitedamagesindicatingscavenging andevenimportationoflionbodypartstotheirdensby LatePleistocenespottedhyenas[ %* ].InnorthernGermany, theonlyknownLatePleistoceneliontrackwayhasbeen documentedwithotherLatePleistocenemegafaunatracks atBottrop,whereadditionallypartlychewedanddamaged lionremainshavebeendescribed[ %* ].FromtheUpperRhine Valley(D)threemoreskeletonremainsandseveralbones havebeenreportedin!"#![ %% ].Here,anewunpublished skullisaddedofalionessfromtheAustrianBadelCave, anotherhyenaandcavebearden,butalsoaskullfromthe AustrianMixnitzCave,orUpperRhineGraenopenairsites, suchasnallythesmallestandmostrarecranialmaterialof siblingandcubremainsfromSwabiancavesofsouthwestern Germany( Figure' ). Additionally,intheLatePleistocene,therewerelarge wolf"ecomorphs"[ )* ]whichglacialsubspecies Canislupus spelaeus Goldfuss,#)!&)[ % ]wasnotyetrevised,asthird largestcarnivores.(eirecologyisstillonlybeginningtobe understood,especiallyfortheircaveoccupation:denusefor cubraisingorimportaspreybyhyenasespeciallyinboreal forestenvironments[ &% $* )* ].AttheSophie'sCave(D), clearevidenceforcavebearscavengingactivitywaspresented

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PaleontologyJournal 10 cm (5a) (5b) (6a) (6b) (8a) (8b) (9a) (9b) (10a) (10b) (12a) (12b) (13a) (13b) (14a) (14b) (15a) (15b) (16a) (11a) (17a) (17b) (18a) (18b) (19a) (19b) (20a) (20b) (21a) (21b) (22a) (22b) (23a) (23b) (36a) (33a) (33b) (34b) (35b) (24a) (24b) (25a) (25b) (7a) (34a) Tooth change (1a) (1b)(2a) (2b) (3a) (4) (3b) Developped saggital crest Non-fused sutures (27b) Chewed (7b) (26a) (26b) (27a) (28b) (28a) (30a) (16b) (11a) (11b) Holotype skull Paratype skull (37b) (37a) (36b) (29a) (29b) (30b) (31) (32) (38) Sligth convex Flat Strong convex Type C Type A Type B Ontogeny Young cub Early adult (always convex) No saggital crest Sligth saggital crest Few used teeth C. c. spelaea No saggital crest Sibling Older cub Fully changed teeth Nearly fully changed teeth (C still growing) (35a) F01234!: Crocutacrocutaspelaea (Goldfuss,#)!&)ontogeneticallyskullshapechangeandthreemainskullshapetypesacrossEurope(composedfrom[ #* ] andnewmaterial).(#)Sibblingskull(withcanniba listicdamage)intoothchangefromSrbskoChlum-Kom nCave,CzechRepublic(NMPNo.R&**/),(a) lateraland(b)dorsal.(! )YoungcubwithlasttoothchangeofthecaninefromUkraine(UZMwithoutno.),(a)lateraland(b)dorsal.(& )Skullofanolde rcub withfullychangedpermanentdentitionfromPerickCaves,Germany(SNSDNo.Sundwig-#' ),(a)lateraland(b)dorsal.($ )Skullofanearlyadultwithf ew usedteethandstillunfusedbraincasesuturesfromBadelCave,Austria(MO MNo.F#),dorsal.(')Skull(withcannibalisticdamage)ofamalefromPeri ck Caves,Germany(BMNHLNo.!)''*),(a)lateraland(b)dorsal.(%)Skull(wit hcannibalisticdamage)fromPerickCaves,Germany(SNSDSundwig-#$),(a ) lateraland(b)dorsal.(* )SkullfromTeufelskammerCave,Germany(RENo.''$*/#A#""'),(a)lateraland(b)dorsal.() )Skull(withcannibalisticda mage)

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% PaleontologyJournal fromErkenbrechtsweileropenairkarstsite,Germany(SMNSNo.#/"%!a),(a)lateralmirroredand(b)dorsal.(/)Earlyadultfemaleskull (withcannibalisticdamage)fromBadWildungen-Biedenstegopenairsite,Germany(HMBWNo.Bi-'!/$'),(a)lateralmirroredand(b) dorsal.(#")SkullofamalefromTrmiceopenairloesssite,CzechRepublic(MB.Ma./"!),(a)lateraland(b)dorsal.(##)Adultskullfrom K onigsaue,Germany(LSDANo.HK-%!:!$)),(a)lateraland(b)dorsa l.(#!)Oldadultfemaleskull(withcannibalisticdamage)fromD obritz Cave,Germany(MBNo.Ma.$$&)#),(a)lateraland(b)dorsal.(#&)Oldadultskull(withcannibalisticdamage)ofafemalefromPerickCaves, Germany(NMBNo.Heinr-#),(a)lateraland(b)dorsal.(#$)AdultskullfromZoolithenCave,Germany(GMBNo.M.!%"/;Holotype inGoldfuss,#)!&),(a)lateraland(b)dorsal.(#')Highadultskull(withcannibalisticdamage)ofafemalefromPerickCaves,Germany (BMNHLNo.!)'')),(a)lateraland(b)dorsal.(#%)Adultskull,withlowerjaw,ofafemalefromSloupCave,CzechRepublic(NHMW No.!"")z"")*/""""),(a)lateralmirroredand(b)dorsal.(#*)Oldadultskullofafemalewithlowerjaw(withcannibalisticdamage)from ZoolithenCave,Germany(UENo.GL**/!"&and!)/'"/;ParatypeinDiedrich,!"##e).(#))Oldadultskull(withcannibalisticdamage) fromSandfortHillCave,England(TMNo.$$/#//'/%/';originaltoReynolds,#/"!),(a)lateraland(b)dorsal.(#/)Oldadultskull(with cannibalisticdamage)fromSandfortHillCave,England(TMNo.$'/#//'/$"*;originaltoReynolds,#/"!),(a)lateraland(b)dorsal.(!") Highadultskullfrom(iedeopenairgypsumkarstsite,Germany(MBNo.Ma.$/#&/.#),(a)lateralmirroredand(b)dorsal.(!#)Highadult skullofamalefromKetsch-KreuzwieseRhineRiverterraceopenairsite,Germany(SMNSNo.%%#*.*.&.%!.#),(a)lateraland(b)dorsal.(!!) Oldadultskull-compositefromBottropopenairsiteandEmscherRiverterraceopenairsite,Germany(MFUOBwithoutNo.),(a)lateral and(b)dorsal.(!&)HighadultskullofafemalefromOaseCave,Romania(SIRBNo.Oasescrocuta-#),(a)lateraland(b)dorsal.(!$)High adultskullfromIrpfelCave,Germany(SMNSNo.AH!%!),(a)lateralmirroredand(b)dorsal.(!')Adultskull(withcannibalisticdamage) fromSloupCave,CzechRepublic(NHMWNo.!"")z "")*/ """!),(a)lateralmirroredand(b)dorsal.(!% )Highadultskullofamalefrom Westeregelngypsumkarstopenairsite(MLU.IFGNo.WEgeln-#&A-B),(a)lateraland(b)dorsal.(!*)AdultskullfromBadelCave,Austria (WMMNo.F!),(a)lateraland(b)dorsal.(!))Deformedhighadultskull(withcannibalisticdamage)ofafemalefromBadWildungenBiedenstegopenairsite,Germany(HMBWNo.Bi-#"at),(a)lateraland(b)dorsal.(!/)HighadultskullofafemalefromGernsheim, Germany(MSGwithoutno.),(a)lateral(mirrored)and(b)dorsal.(&")HighadultskullfromDoesburgnearArnheim,Germany(RE withoutno.),(a)lateral(mirrored)and(b)dorsal.(&#)YoungadultskullfromPinHoleCave,England(MMUNo.%'/*),lateralmirrored. ( &!)AdultskullfromtheMlade cCave,CzechRepublic(AMBNo.!%$$),lateral.(&&)YoungadultskullofafemalefromSrbskoChlumKom nCave,CzechRepublic(NMPNo.R#"%*),(a)lateraland(b)dorsal.(&$)OldadultskullfromthefemaleskeletonofV ypustekCave, CzechRepublic(NHMWNo.A''!/),(a)lateralmirroredand(b)dorsal.(&')Adultskull(withcannibalisticdamage)fromtheJavorkaCave, CzechRepublic(AMBNo.OK##$)/#),(a)lateraland(b)dorsal.(&%)Highadultskull(withfrontalbiteimpactintheprocessofhealing) fromIrpfelCave,Germany(SMNSNo.*.)"#),(a)lateraland(b)dorsal.(&*)HighadultskullofamalefromGernsheim,Germany(MSG withoutno.),(a)lateral(mirrored)and(b)dorsal.(&))Senileskullwith 4 dentalpathologyfromCrumstadt,Germany(HLMDwithout no),lateral. basedonbonefragmentsfoundinexcrementsandwithin afaecalplacewherewolvesevenusedapartofacavebear den(severalmetersdeepfromtheentrancearea)overashort periodobviouslyasden[ $* ]. 2.MaterialsandMethods &"&hyenaand#. &*&lionbonesofLatePleistoceneage from#"%studiedcaveandopenairlocalitiesinCentral Europewereanalysedmainlyfrompreyboneassemblages ofseveralGermanandCzechhyenadens( Table# ).( is studyandreviewusedaninterdisciplinaryapproachapart fromclassical"bitemarkanalysesonsinglebones",rather itusedthenewlydeveloped"butcheringdecomposition andbonedamagestage"analyses.Newresultsstartedwith therediscoveryofmanyhistoricallycollectedbonesofthe LatePleistocenepredatorsandpreyremains,whichwere reidenti-edforthe-rsttimeashyenadenorigin.Hyena populationswereanalysed,alongwiththeboneassemblages. (estudyofallanimalgroupsbasedonNISP(=number ofidenti-edspecimenspertaxon)andpartialMNI(= minimumnumberofindividuals)analyseswerealsothebasis todistinguishdensitetypesusingthemodernhyena/lion ethologycomparison.Importantfortheresultswasthestudy ofdi+erenttopographicrelatedhyenadenmorphotypes: cavesopenair,riverterrace,loess,andgypsumkarstsites. ( ehistoricalcollectionsarehighlyusefulfortheseanalyses, andinseveralcasestwothingswereimportantfortheirmodernanalysesuse:(#)bonesfromdi+erentcollectionswere compiledfromdi+erentmuseums;thiso.endemonstrated theanthropogenicselectionof"goodbones"and"badbones" (bonefragments),butinhyenadenanalysesthecomplete assemblageisrequiredforinterpretations.Insomecases thehistoricalsiteswerecompletelyrestudiedattheexisting sitessuchastheZoolithenCave,Sophie'sCave,Srbsko Chlum-Kom nCave,SloupCave,Westeregelnopenair,and Sewecken-Bergeopenair,whereadditionalexcavationsand stratigraphic/sedimentologicalandC#$datingworkwere importanttounderstandtheage,bonetaphonomy,and sitegenesisingeneral.Intotal,severaltensofthousands ofboneswereanalysedsince!""$fromthreemainstudy areas(SauerlandKarst,(uringianKarst,Bohemainand fewerMoravianKarst,FranconianKarst,smallareainthe CarpathianApuseniMountainkarst,andopenairsitesin Germany/CzechRepublic).Aspartofthosestudiesthe holotypeskullsof C.c.spelaea P.l.spelaea ,and U.s. spelaeus wererediscoveredinthefamousRosenm uller#*/$ collection.OtherforgottenrediscoveredGermanhyenaden sitecollectionsarefromGiebel#)$$#)$)(Westeregeln, Sewecken-Berge),Nehring#)*!( (iede,Westeregeln),or M uller#/"!(FuchslukenCavity).(islargeamountofmaterialstudiedisnomoreattheformerPreu§ischeGeologische LandesanstaltBerlin(nowcoll.MB).Additionally,allthe mostlyoverlookedindividualskeletonsofhyenas/lionshave beenidenti-ed,prepared,andcomposedbytheauthorinthe pastyearsinvariouscollections.(emainimportantand relevantsitesarelisted( Table# )atwhichmaterialishoused inthefollowinginstitutionsofdi+erentcountries:Austria:

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PaleontologyJournal Anterior Posterior E Round F Irregular G Drop 5 cm Type C Oval Type B Disk Type A Conical Type G Drop Type F Irregular Type D Long-oval C a a b b (1a) Broken cusp (polished roots) Lost tooth (1d) Broken cusp (polished roots) P 3 P 4 M 1 P 3 P 4 M 1 P 2 M 1 P 3 P 2 C Broken cusp (polished roots) P 3 P 4 P 2 (1c) (1b) (2a) (2b) P 3 P 3 P 4 P 3 Lost tooth P 4 (2c) (2d) Broken cusp (polished roots) Lost teeth, fused alveols, polished root Lost tooth, fused alveols Broken cusp (polished roots) Lost teeth P 3 P 4 P 2 M 1 M 1 Lost teeth, fused alveols Half broken tooth Half broken tooth P 4 Lost tooth, fused alveolus Half broken cusp (polished root) P 2 (3a) (3b) (3c) (7a) P 3 P 3 P 4 P 4 P 2 M 1 (6) (7b) (8a) (8b) (9) (10) Type E Round Pellet shape types Lost tooth and partly closed alveols (4a) (4b) Lost tooth and closed alveols (5a) (5b) P 2 P 3 P 4 M 1 (11) (12) Broken cusp (polished roots) Lost tooth and closed alveols Displaced tooth Broken cusp (polished roots) P 4 P 4 P 3 P 2 P 1 P 4 P 3 P 2 P 1 P 1 P 1 P 4 P 1 P 1 P 2 P 4 P 3 P 2 P P 4 P 3 P 2 P 1 I 1 I 2 I 2 I C C All I C 3 1 Bone crushing dental pathologies cracked 10 cm a b Adult skeleton (cracked) Humerus Ulna Femur Tibia Radius Atlas Skull 10 cm (a) (c) (b) A Conical B Disk C Oval D Long-oval Srbsko Chlum-KomÂ’n Cave (CZ) Bone F01234&:(a) Crocutacrocutaspelaea (Goldfuss,#)!&)dentalpathologiesonskullsandlowerjaws.Maindamagesarefoundonthepremolars (bonecrushing)andtheincisives/canines(nibbling)(composeda.er[ '/ ]).(#)ParatypeskullofanadultwithlowerjawfromtheZoolithen Cave,Germany(UENo.GL**/ !"&and!)/ '"/),(a)lateral,(b)ventral,(c)lowerjawdorsal,and(d)lowerjawlateral.(! )Skullwithlower jawofasenilefromPerickCaves,Germany(NMBNo.Heinr-#),(a)lateral,(b)ventral,(c)lowerjawdorsal,and(d)lowerjawlateral.(&) SkullofanoldadultfromSandfortHillCave,England(TMNo.$$/#//'/%/';originaltoReynolds,#/"!),(a)lateral,(b)ventral,and(c)lower jawdorsal.($)SenileskullfromCrumstadt,Germany(HLMDwithoutno),(a)lateraland(b)ventral.(')HighadultskullfromGernsheim, Germany(MSGwithoutno.),(a)lateral(mirrored)and(b)ventral.(%)HighadultrightmandiblefromthePerickCaves,Germany(GMB no.M!'%'),lateral.(*)Senilele.mandiblefromtheWilhelmsCave,Germany(GMBno.M'///(Mandible)/GPIMno.A'F#!)#(M#),(a) lateraland(b)dorsal.() )HighadultmandiblefromtheSewecken-bergeopenairgypsumkarst,Germany(MBno.Ma!/%"%),(a)dorsal and(b)lateral.(/)HighadultrightmandiblefromtheGr urmann'sCave,Germany(GMBwithoutno.),lateral.(#" )Senilerigthmandible fromthePerickCaves,Germany(SNSDno.Sundwig-$$),lateral.(##)Highadultle.mandiblefromtheWilhelmsCave,Germany(EMSCH No.Frettertal-#),lateral.(#!)AdultmaxillaryandmandiblefromtheReporjeKalvarieCave,CzechRepublic(NMPno.'$*&/R#!and###), lateral.(b)ByIceAgespottedhyenascannibalisticscavengedandcrackedhyenacarcassfromtheSrbskoChlum-Kom nCavehyenaden, CzechRepublic(modieda.er[ $# ]).(c)IceAgespottedhyenacoprolitetypeshapetypes(modieda.er[ %" ]).

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) PaleontologyJournal Panthera leo atrox Panthera leo spelaea Panthera leo subsp. Late Pleistocene (113,000-24,000 yBP) Panthera leo tigris (a) Srbsko Chlum-Kom’n Cave (CZ) Neumark-Nord Lake 1 (D) (d) Panthera leo vereshchagini London Berlin Paris Warsaw Oslo Moskow Bukarest Rome Athene Barcelona Madrid Munich Amsterdam Budapest Glacier Lions in alpine regions Glaciers Panthera leo spelaea bones Tundra Forest tundra Boreal conifere forests Temperate broadleaf forests Mixed forests Taiga steppe Mediterranean forests Mountains with few vegetation Skeletons, skeleton remains Zoolithen Cave (D) Perick Caves (D) Westeregeln (D) Sloup Cave (CZ) V !pustek Cave (CZ) Ursilor Cave (Ro) (b) Hutton Cave (GB) Wookey Hole Cave (GB) Bleadon Cave (GB) Townton Cave (GB) Agios Georgios Cave (Gr) Sophie's Cave (D) Sandfort Hill Cave (GB) Pin Hole Cave (GB) Medvedia Cave (Sk) iede (D) Badel Cave (A) 500 km Carpathains Dianrids London Berlin Paris Warsaw Oslo Bukarest Athene Barcelona Madrid Munich Amsterdam Budapest Alps Carpathains Dianrids Alps Glacier Hermann's Cave (D) Sarajevo Perick Caves (D) Sloup Cave (CZ) Zoolithen Cave (D) Sophie's Cave (D) Wei§e Kuhle Cave (D) Ursilor Cave (Ro) M o s t n o r t h e r n c a v e b e a r s Keppler Cave (D) M o s t n o r t h e r n l i o n s Cave bears in mountain regions Rome Late Pleistocene (MIS 3-5d) Mixnitz Cave (Au) Gro§e Teufels Cave (D) Balve Cave (D) V !pustek Cave (CZ) Genther Cave (D) Sybillen Cave (D) Medevieda Cave (Sk) Badel Cave (Au) Pruzinsk‡ Cave (Sk) Ice Age Stepe lion skeletons in cave bear dens Cave bear dens Ice Age steppe lion bones in cave bear dens Mixnitz Cave (A) Wildkirchli Cave (Ch) North Sea (c) Arrikrutz Cave (Es) AzŽ Cave (F) Santenay (F) VerzŽ Cave (F) Siegsdorf (D) Huttenheim (D) Srbsko Cave (CZ) Komarowa Cave (Pl) Trmice (CZ) Deszczowa Cave (Pl) Neumark Nord I (D) Bottrop (D) Beringia F01234$:IceAgesteppelion Pantheraleospelaea (Goldfuss,#)#")distribution(a)worldwideand(b)importantsitesincentralEuropeduring Early/MiddleLatePleistocenebeforeLGM(##&."""!$."""BP)(composedfromdi+erentsources;seereferencelist).(c)Lionskeletonsand remainsinLatePleistocenecavebeardensofEurope.(d)ArticulateddiseasedlionessskeletonfromNeumark-NordLakeI(D)elephant graveyard(from[ %# ])andsubadultdiseasedlionessfromSrbskoChlum-Kom nCavehyenaden(CZ)(from[ %! ]).

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PaleontologyJournal / 1e Developped saggital crest Non-fused sutures Non-fused sutures 10 cm Holotype skull P 4 M 1 C P 4 P 3 dm 4 dm 4 dm 3 Tooth change (2b) (4) (2a) (3) (5) (6a) (6b) (17a) (18a) (22a) (22b) (8a)(8b)(9a)(9b) (10a) (10b) (11b) (12a) (12b) (13a) (13b) (14a) (14b) (15a) (16a) (19a) (20a) (21a) (23b) (23a) (17b) (18b) (15b) (16b) (19b) (20b) (21b) dm 4 dm 3 (1a) (1b) (11a) (7a) (7b) Ontogeny P. l. spelaea Cub Early adult Sibbling No saggital crest ( 17a ) ( 18a ) F01234': Pantheraleospelaea (Goldfuss,#)#")ontogeneticallyskullshapechangeandthreemainskullshapetypesacrossEurope(composed from[ ) $% %! %% ]andhereinaddednewmaterial:#&,#", #&, #%and!").(#)SibblinglowerjawfromtheMixnitzCavecavebearden,Austria (REno.NMB#")a),(a)dorsaland(b)lateral.(!)IncompletesiblingskullfromAu5auserCavehyenaden,Germany(SMNSwithoutno.), (a)dorsaland(b)lateral.(&)SiblingmandiblewithmilkdentitionfromHohlensteinCavehyenaden,Germany(SMNSno.&&!!%),lateral. ( $ )CubmaxillarywithpermanentdentitionfromZoolithenCavehyenaandcavebearden,Germany(UEno.Sp$!/ #$),lateral.(' )Cub mandiblewithpermanentdentitionfromZoolithenCavehyenaandcavebearden,Germany(UEno.GL**/)'),lateral.(%)Earlyadultskull fromaskeletonofUrs ilorCavecavebearden,Romania(SIERno.PU/"""#),(a)dorsaland(b)lateral.(*)Earlyadultskullwithlowerjaw fromaskeletonofSrbskoChlum-Kom nCavehyenaden,CzechRepublic(NMPno.R$$"%),(a)dorsaland(b)lateral.() )Earlyadultskull fromthePerickCaveshyenaandcavebearden,Germany(BMNHLno.!)''&),(a)dorsaland(b)lateral.(/)EarlyadultskullfromBerounH yskovopenairriverterracesite,CzechRepublic(MBKBno.&%&a),(a)dorsaland(b)lateral.(#" )AdultskullfromZoolithenCavehyena andcavebearden,Germany(MBno.Ma.'"/$*),(a)dorsaland(b)lateral.(##)AdultskullfromBadelCavehyenaandcavebearden,Austira (MOMno.F# ),(a)dorsaland(b)lateral.(#! )AdultskullfromSloupCavehyenaandcavebearden,CzechRepublic(AMBno.OK#&"'*"), (a)dorsaland(b)lateral.(#&)SenileskullfromCampusalCave,Romania(SIERwithoutno.),(a)dorsaland(b)lateral.(#$)Adultskullfrom

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#" PaleontologyJournal Bobenheim-Roxheimopenairriverterrace,Germany(REno.PCC#&!),(a)dorsaland(b)lateral.(#')AdultskullfromZoolithenCave hyenaandcavebearden,Germany(UM-Ono.BT'$!#),(a)dorsaland(b)lateral.(#%)Senileskullwithseveraldentalpathologiesfrom Hermann'sCavecavebearden,Germany(NMBno.#*/$-M),(a)dorsaland(b)lateral.(#*)Adultskull(mountedincompositeskeleton) fromSloupCavehyenaandcavebearden,CzechRepublic(AMBwithoutno.),(a)dorsaland(b)lateral.(#))Adultskull(mountedin compositeskeleton)fromSloupCavehyenaandcavebearden,CzechRepublic(NHMVno.#))'/""#$/$&"!),(a)dorsaland(b)lateral.(#/) AdultskullofaskeletonfromHuttenheimopenairriverterracesite,Germany(SMNSno.%)#%.'.%.*&.#),(a)dorsaland(b)lateral.(!")Adult skullofaskeletonfromEdingen(Br uhl)openairriverterracesite,Germany(SMNSno.%%#*. # /. *!. ),(a)dorsaland(b)lateral.(!# )Adult skullfromMixnitzCavecavebearden,Austria(REno.NMB#"*),(a)dorsaland(b)lateral.(!!)Holotypeskullwithbitemarkdamage pathologyofanadultmalefromZoolithenCave,Germany(MBno.Ma.'"/$)),(a)dorsaland(b)lateral.(!&)AdultskullfromZoolithen Cavehyenaandcavebearden,Germany(MBno.Ma.$)#''.#),(a)dorsaland(b)lateral. Cracked Lower jaw removement Braincase opening Cracked Bite damages Cracked Cracked Bite impacts Bite impacts 10 cm Cracked Bite impacts, scratches Cracked Cracked Cracked (2) (3) (1) (4) (5) (6) (7) (8) Cracked (a) Chewed 4 P 4 C 10 cm I 1 Modern damage Bite marks Ulna Metapod Humerus Canine Skull Mandible P4 tooth Metapod Lumbar vertebra Metapod Metapod Metapod Metapod Metapod Femur Lumbar vertebra Radius Mandible Phalanx 1 Phalanx 1 Phalanx 2 Ulna Phalanx 1 Phalanx 2 Lumbar vertebra Cave bear nests Adult female cave bear skeleton Cave bear, one year old cub skeleton 100 cm Cave wall N Humerus Scattered and scavenged young lioness ?Mother and cub? A Cub Se 100 m Cave bear nests (more then 140) River Exit Entrance Dropped blocks Speleothems End of Visitor loup Trail Lion skeleton 1 ? N Female cave bear and cub skeletons Male cave bear skeleton 4 Lion skeletons (articulated) Cave bear skeletons (articulated) Ursilor Cave (Ro) Hyena den area Lion skeleton 2 Lion skeleton 3 Lion skeleton 3 Lion skeleton 1 Lion skeleton 4 N Lion skeleton 3 Skull P4 tooth Radius Ulna Ulna Humerus Canine Metatarsalia Metatacarpalia Phalanx 1-2 Lumbar vertebra Humerus Skull Scapula Ulna Tibia Femur Pes Pes Manus Manus Costae Vertebrae Pelvic Collapse cone 10 cm Cracked Chewed Cracked Chewed Chewed Chewed (9) (10) (11) Chewed (12) Cracked (13) (14) (17) Chewed Chewed Chewed Chewed (18) (19) (20) (21) Chewed (22) (23) Bite scratches Chewed Chewed Chewed Chewed (16) Chewed (15) Chewed Bite marks (24) (25) (b) 50 cm F01234%:IceAgesteppelionpostmortalbitedamageson(a)cranialandpostcranialbonesand(b)skeletonofasubadult(scavengedand scatteredbesidesmotherandcubonplaceskeletonsonhibernationplateau)deepintheUrs ilorCave(Ro)(composedfrom[ %' %* ]).

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PaleontologyJournal ## Bite tr auma !" cm Bite trauma Bite trauma B ite trau ma Bite trauma B ite trau ma deforma tion Exostoses Exostoses ?Bite damage Exostoses Exostoses Fusion Fusion (Exostoses) Broken bula Fusion BrŸhlKoller (D) Neumark-Nord Lake (D) Zoolithen Cave (D) Exostoses Exostoses Broken bula Broken bula Fusion Exostoses Zoolithen Cave (D) Sloup Cave (CZ) B ite traum a deformat ion Crushing Catching P # P # P # P $ P $ P % I $-! I $ Catching Nibbling Cutting F01234*:(a)IceAgesteppelionand(b)IceAgespottedhyenaalivetraumapathologies(bitedamagesmainly)fromdi+erentsitescaused byhyenas,lions,andcavebearsduetobitedamagesinbattlesinandoutsideofcaves(compiledpartlyfrom[ '/ %) ]).

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#! PaleontologyJournal Cave dwellers Cave periodic occupants 99% 6% Hyena den Cave bear beds Hibernation area Cave bears Small cave branch Cave bear den More secure zone Conflict zone "Vertical shaft" Hyena antagonist conflicts Juvenile AdultOlder 100 Individual ages Adult 0 Ursus spelaeus spelaeus and ingressus Panthera leo spelaea Crocuta crocuta spelaea Older Juvenile Cub Mortality rate 16% 56% 28% Cub Juvenile AdultOlder NISP = 2.600 80% 16% 4% NISP = 206 AdultOlder Cub Juvenile NISP = 229 Lions stealing hyena prey, killed in cave by hyenas 10 cm Ursus Partly healed bite damage Panthera Crocuta Partly healed bite damage Partly healed bite damage Lions killed by cave bears during predation on cubs and hibernationg bears (a) (b) (c)(d) F01234):LatePleistocenespottedhyenas,steppelions,andcavebearethology(cavedwellersorinhabitants)andcon,ictmodelforlarger cavesystems(hereZoolithenCave,composedfrom[ $" $# ],cave-imagingillustrationsG.Teichmann).

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PaleontologyJournal #& Canis lupus subsp. Panthera leo spelaea Crocuta crocuta spelaea Putorius eversmanni Capra ibex Coelodonta antiquitatis Bison priscus Mammuthus primigenius Equus caballus przewalskii Megaloceros giganteus Cervus elaphus Rangifer tarandus Equus caballus hydruntinus Gulo gulo Panthera pardus Alopex alopex Vulpes vulpes Martes martes Cave bear predation/scavenging specialization by absence of steppe fauna Lepus timidus/ europaeus Meles meles Ursus cf spelaeus Rupicapra rupicapra Westeregeln NISP = 572 Sewecken-Berge NISP = 775 Mammoth Steppe Fauna 1% 22% 5% 1% 7% 10% 4% 37% 11% 2% Teufelskammer Cave NISP = 199 Perick Caves NISP = 2.275 Boreal forest cave bear Fauna Bottrop NISP = 3.820 Mixed Fauna Bilstein Cave NISP = 1.120 1% 8% 12% 1% 67% 4% 8% 1% 4% 3% 1% 0,1% Near absence of Cave bear Absence of Mammoth 1% 1% 94% 2% 0,1% 2% Mammoth steppe Boreal Forests Mountains River valleys/ mountain margins (Saiga, Ovibos) (Ursus) (b) Bad Wildungen NISP = 260 Hyena/Cave bear den Hyena/Cave bear den Cave bear den 2% 32% 5% 8% 8% 10% 2% 5% 2% 1% 11% 10% Hyena/Fox/Badger den 1% 2% 15% 2% 25% 0,1% 11% 2% 6.7% 0,1% 3% 0,1% 1% 0,1% 34% Hyena den Gypsum karst 14% 5% 26% 1% 19% 15% 0,1% 1% 0,1% 5% 2% 10% 1% 2% 0,1% Hyena den Westeregeln (D) n = 84 1% 97% Bottrop (D) n = 50 10% 12% 78% Prey depot den Birth den Commuting den Single hyenas Hyena clan Hyena mother with cubs 2% Bad Wildungen (D) n = 7 Roter Berg Saalfeld (D) n = 70 43% 57% 36% 7% 57% Sewecken-Berge (D) n = 114 4% 94% 2% (a) WŸste Scheuer Abri (D) n = 27 4% 30% 66% Ice Age spotted hyena den types (after ethology) Cave den Open air karst den Open air river terrace/steppe den Hyena den Hyena Lion conflicts (after morphology/landscape) (c) 0,5% 17% 3% 2% 0,5% 44% 0,5% 5% 20% 1% 4% 1% 1% 0,5% 0,5% 0,5% Hyena den (d) (e) Mammoth steppe fauna prey Sinkhole Prey Sibbling Cub Adut to senile Ice Age Spotted hyena Ice Age steppe lion Ice Age wolf Horse/bison hunting specialization by rarenes of cave bears Mammoth Cave bear Woolly rhinoceros Przewalski horse Ice Age donkey Steppe bison Musk ox Giant deer Red deer Reindeer Saiga antelope Der antler collectors Boreal forest fauna prey F01234/:(a)IceAgespottedhyenadentypesa.erethologyandlandscapeandmorphology.(b)Hyenapreyboneassemblagesand(c)prey specializationinmammothsteppelowlandandborealforestmountains.(d)IceAgespottedhyenasdecomposinginbutchertechniquea woollyrhinoceroscarcass.(e)IceAgespottedhyenaclanhunti ngPrzewalskihorse(illustrationsG."Rinaldino"Teichmann).

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#$ PaleontologyJournal NHMV=NaturalHistoryMuseumVienna;CzechRepublic: AMB=AnthroposmuseumBrno,MBKB=Museumofthe BohemianKarstBeroun,andNMP=NationalMuseum Prague;England:BMNHL=BritishMuseumofNatural HistoryLondon,England,SM=SomersetMuseum;Germany:BM=BalveMuseum,GMB=Goldfussmuseum Bonn,GPIM=Geological-PaleontologicalMuseumofthe WestphalianWilhelmsUniversityM unster,GZG=GeolocicalCentrumG ottingen,HC=HeinrichsCaveHemer, LDA=LandesmuseumforArchaeologySaxony-Anhalt, LM=L obbeckeMuseumAquazooD usseldorf,ME=privatecollectionMenger,MLUIFG=Matin-Luther-University InstituteforGeosciencesHalle/Saale,MMS=MammutMuseumSiegsdorf,MNB=MuseumforNatureandHumans Bielefeld,MUOB=MuseumPre-andLocalHistoryBottrop,MSG=MuseumSch o +erstadtGernsheim,RE=privatecollectionReiss,SNMB=StaatlicheNaturhistorische MuseumBrunswig,SMNS=StaatlicheMuseumNaturkunde Stuttgart,SNSD=StaatlicheNaturhistorischeSammlungen Dresden,UE=UniversityErlangen,UZM=UrzeitmuseumTau6irchen,U-MO=Oberfr ankischeUrweltmuseum Bayreuth,andZO=ForschungsgruppeH ohleundKarst Frankene.V.,N urnberg;Romania:USC=Urs ilorCave,SIER =SpeleologicalInstituteEmilRacvitaBucharest;USA:MOM =MuseumofMan. 3.LatePleistoceneSpottedHyenas ( ehyenasoftheEuropeanLatePleistocenewerepopularisedhistoricallyas"cavehyenas"withthelatinbinomial name" Hyaenaspelaea ,"becauseoftheirinitialdiscoveriesin theZoolithenCave(D)[ % ]andothercavesinEurope.Recent revisionaccordingtomoderntaxonomic[ )/ ]andDNA[ /" ] studieshasconsideredtheseLatePleistocenespottedhyenas as Crocutacrocutaspelaea (Goldfuss,#)!&)[ $' ].( eseLate Pleistocenehyenasareslightlydi+eredtomodernspotted hyenasubspecies,withthoseintheEemianinterglacialbeing smallerandtheWeichselianglaciallarger[ /# /! ].Similarto modernspottedhyenas,thefemalesintheextinctspecies[ /& ] arelarger[ $' /$ ],whichisalsore, ectedintheskullrecord ofEuropefortheLatePleistocenespottedhyenas( Figure! ) [ #* ]. & .HyenaPopulationsoverEurope. Inthestudyareaof GermanyandCzechRepublic,theLatePleistocenefossil recordincludes!,&"&bonesand!skeletonsfromhyenas ( Table# ).( esepredatorsmusthavemigratedfromAfrica totheNorthandweredistributedalloverEurasiaupto northernEngland(mostnortherndeninPitHoleCavern (GB))duringtheLatePleistocene(Eemianinterglacialto Weichselian/Wuermianglacial,#!%."""!$."""BP).(elast hyenasofEuropemusthavebecomeextinctjustbeforethe LastGlacialMaximum(=LGM,around!!,"""BP)[ /& ]at leastinnorthernEurope.Aretreattothesouthisnotreally expected,whileaneasternmigrationisnotyetsupported bystudiesfromAsiawithmosteasternrecordsintheAltai Mountains.Inmostcasesexceptinthewellstudiedregions ofGermanyandCzechRepublic,wherehyenaremainsare found,theyoccuratdensiteseventhoughtheyhavebeen mostlyoverlooked( Figure# ).Anoverviewofmorethan#!" LatePleistocenehyenadensites( Figure# )indicatesmuch moresmallcavesandentrancesoflargercavestohavebeen used,whereastherareopenairrecordismostlyaresult ofbonetaphonomyaccumulationmisidenti-cation.Notall sitesaredatedproperly,buttheylikelyrepresenttheLate Pleistoceneinterval. '.#.Holotype,ParatypeSkulls,andSkeletons. ( erecently rediscoveredholotypeskullof" Hyeanaspelaea "wasfoundin theZoolithenCaveofBavaria(D)[ % ].Besidesthistoothless, oldindividual,andhistoricallydamagedskull,aparatype skullwasselectedfromalargehyenapalaeopopulationof thiscave[ $$ ].Fromthiscave,similartomanyothersitesof Europe,articulatedskeletonsareabsentduetotworeasons: (a)nonprofessionalexcavationandcollectingand(b)hyena cannibalism.Morerecently,afairlycompleteindividual skeletonofanadultanimalwasrediscoveredandcomposed fromtheKon eprusyCave(CZ)[ $" ]( Figure# ).Another skeletonwithabetterpreservedskullwaschosenfrom theV ypustekCave(CZ)[ $# ]( Figure# ).Skeltonremainsof threedi+erentcub/siblingindividualswereusedfromthe Srbsko-Chlum-Kom nCave[ $# ],tobuilda-rstcomposite cubskeletonofthisspecies( Figure# ),whichisimportant fortheidenti-cationofoverlookedcubbonesinvarious collections.Furtherincompletehyenaskeletonsseemtohave beenpresentalsointhePerickCaves,HuttonCavern,and ZoolithenCave,butcannibalism,river,oods,orhistorical excavationsdestroyedtheirarticulationcontext. ' .SkullShapeTypes. Recentstudiesonboth,modern Africanspottedhyenas[ /$ ]andmorethan&'European LatePleistocenespottedhyenaskulls(mainlygrownup individuals,butalsofewsiblingandcubskulls)[ #* ],have yieldedsimilarresults.( esaggitalcrestisabsentininfants anddevelopswithinjuvenilesa.ertheirteethchangeandis stronglyhighinadults( Figure! ),whichservesastheattachmentforthemassivebonecrushinglowerjawmusculature. ( reedi+erentgeneralskullshapevariantssimilarin C.c. spelaea andin C.c.crocuta include(a), atand(b)slight convexsaggitalcrestshapes( Figure! ).Asdemonstratedin EuropefortheLatePleistocenespottedhyenas,thosetypes overlapanddonotallowaclearseparationof"races"(Figures # (c)and ).( ethird,(c)strongconvexshape(only'%of theskulls),wasdemonstratedfortheLatePleistoceneand modernspottedhyenastoberareandpathologicandpartly theresultofcranialbitedamagescausedbytoppredators incon, icts(otherhyenasorlionsorcavebears;seeFigures and )[ #* ].(ebestandmostimpressivedamagedskull isfromtheZoolithenCave( Figure* (b)),whichhistorically [ /! ]astonishedscientists,becausethebraincasewasnot damaged,andtheindividualsurvivedatleastcouplesof dayswithstrongskullbitetraumadamage(twocentimeter deepsaggitalcrestdamage, Figure* (b)).Fullhealingofsuch traumasresultedinhigh-convexsaggitalcrestshapessimilar tothosefoundattheopenairsiteofBr uhl-Spielwiesen(D) ( Figure* (b))[ $) ].

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PaleontologyJournal #' T7894#:!.&"&hyenaand#.&*&lionbonesofLatePleistoceneagefrom#"'studiedcaveandopenairlocalitiesinCentralEuropewithIce Agespottedhyenaandsteppelionremains(detailsaboutthematerial,localitypositions,andhousingincollectionscanbefoundinthe references). LocalitySitetypeAge Hyenas (NISP) Lions (NISP) Bone assemblage (NISP) References Ahlen(D) Openairloess,mammoth skeletonscavengingsite Weichselian (glacial) Indirectby scavenging marks Mammoth skeleton scavengingsite [ '' ] Al .er(D) Openairloessonriver terrace Weichselian (glacial) "!?unclear[ %# ] Altlussheim, Almendwiesen (D) Openairloessonriver terrace Weichselian (glacial) ##'%![ %# ] Altlussheim, Eichelgarten(D) Openairloessonriver terrace Weichselian (glacial) "#?unclear[ %# ] Altlussheim, Silzwiesen(D) Openairloessonriver terrace Weichselian (glacial) "#?unclear[ %# ] Altrip, Neuhofener Altrhein(D) Openairloessonriver terrace Weichselian (glacial) "#?unclear[ %# ] BadK ostritz(D) Zechsteingypsumkarst hyenaden (preydepotandcommunal den) Weichselian (glacial) !$?unclear[ %# ] BadK osen(D)Openairloess Weichselian (glacial) "#?unclear[ %# ] BadLauchst adt (D) Openairloess Weichselian (glacial) % (fromanadult female individual skeleton) ?unclear[ %# ] BadWildungen (D) Openairloessonriver terrace Weichselian (glacial) ##(and#' coprolites) "!&&[ $/ ] BalveCave(D) Carboniferouslimestone cave(cavebearden,hyena cubraisingandcommunal den/overlapping Neanderthal,and Aurignaciancampsite) EemianWeichselian (interglacial, mainlyglacial) &$'%(!skulls)?unclear[ $! %' ] Baumann'sCave (D) Carboniferouslimestone cave(cavebearden,short timehyenaden,? temporarywolfden) ?SaalianEemian, mainlyWeichselian (interglacial, mainlyglacial) #"#$) ?unclear,half Millioncave bearbones unpublished Berlin-Kreuzberg (D) Openairriverterrace Weichselian (glacial) "#?unclear[ %# ] Beroun(CZ)Openairriverterrace Weichselian (glacial) "#(skull)?unclear[ )" ] BilsteinCave(D) Carboniferouslimestone cave(cavebearden,short timehyenaden,? temporarywolfden) Weichselian (glacial) !&/ ?unclear,many cavebear remains [ )! ] Br uhl(Koller), Schlangenwinkel (D) Openairriverterrace Weichselian (glacial) #!bonesofa maleskeleton ?unclear[ )' ]

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#% PaleontologyJournal T7894#:Continued. LocalitySitetypeAge Hyenas (NISP) Lions (NISP) Bone assemblage (NISP) References Br uhl, Spieswiesen-Ost (D) Openairriverterrace Weichselian (glacial) "#?unclear[ %# ] Br uhl, Spieswiesen-West (D) Openairriverterrace Weichselian (glacial) ) (!skulls)$#.!''[ %# ] Br uhl(Koller), Rheingewann(D) Openairriverterrace Weichselian (glacial) #&'##[ %# ] Burgtonna(D)Travertin Eemian (interglacial) # (unclear amount) & (unclear amount) ?unclear[ %# ] Bochum(D)?Openairriverterrace ?Weichselian (glacial) "#(skull)?unclear[ %# ] Bottrop(D) Openairriverterrace (hyenacommunalden, partlycubraisingdenand preydepot) Weichselian (glacial) '" !%(andone trackway) ?,about&.""" bones,manyof hyenaden origin (especially woolly rhinoceros) [ $) %# ] DeutmeckerCave (D) Carboniferouslimestone cave(hyenaden) Weichselian (glacial) !"?unclear[ %' ] Edingen(Br uhl), EdingerRied(D) Openairriverterrace Weichselian (glacial) (skulls) and) bonesof amaleskeleton, includingskull # ###[ )' ] Fl orsheim(D) Openairloessonriver terrace Weichselian (glacial) #"?unclearunpublished Fuchsluken CavitiesSaalfeld (D) Zechsteinlimestonekarst cavitieshyenaden(cub raisingandcommunalden) EemianWeichselian (interglacial, mainlyglacial) !#)/#. "&'[ %# ] Freiburga.d.U. (D) Openairloessonriver terrace,mostprobably MiddlePalaeolithicsite Weichselian (glacial) #&(froman adolescent individual skeleton) ?[ %& ] Geddin(D)Openairriverterrace Weichselian (glacial) "#?unclear[ %# ] GentherCave(D) UpperJurassicdolomite cave(cavebearden) Weichselian (glacial) "#(skull)?unclear[ *# ] G ottingen(D)Openairriverterrace Weichselian (glacial) "#?unclear[ %# ] Gr urmannsCave (D) Carboniferouslimestone cave(hyenaden) Weichselian (glacial) ###?unclear[ %' ] Halle/Saale(D)Openairloess Weichselian (glacial) "#?unclear[ '" ] Haltern(D)Openairriverterrace Weichselian (glacial) "#?unclear[ %# ] Heddesheim, Neuwiesen(D) Openairriverterrace Weichselian (glacial) "!?unclear[ %# ] Hermann'sCave (D) Carboniferouslimestone cave(cavebearden) Weichselian (glacial) $*(oneskull, twoindividual skeleton remains) ?unclear,many cavebearbones [ *% ]

PAGE 17

PaleontologyJournal #* T7894#:Continued. LocalitySitetypeAge Hyenas (NISP) Lions (NISP) Bone assemblage (NISP) References Herne(D)Openairriverterrace Weichselian (glacial) #"?unclear[ $* ] HertenStuckenbusch (D) Openairriverterrace Weichselian (glacial) "!?unclear[ %# ] Hohenm olsen (D) Openairloess Weichselian (glacial) #(skull)"?unclear[ '" ] HohleSteinCave (D) Carboniferouslimestone cave(hyenapreydepot) Weichselian (glacial) ##&'[ %' ] Holede cnear Zatec(CZ) ? Weichselian (glacial) "!?unclear[ )" ] Host m(CZ)Openairriverterrace Weichselian (glacial) "$?unclear[ )" ] HuttenheimHuttenheimer Kammer(D) Openairriverterrace Weichselian (glacial) & and&*bones ofonelioness individual skeleton includingskull ?unclear[ )' ] Huttenheim, Sandfeld(D) Openairriverterrace Weichselian (glacial) "#?unclear[ %# ] JohannesCave (D) Carboniferouslimestone cave(hyenapreydepot) Weichselian (glacial) &!&#[ %' ] Karlsruhe, Neureut(D) Openairriverterrace Weichselian (glacial) "#?unclear[ %# ] KepplerCave(D) Carboniferouslimestone cave(cavebearden) Weichselian (glacial) "&& ?unclear,many cavebear remains [ %' ] Ketsch, Kreuzwiese(D) Openairriverterrace Weichselian (glacial) (skulls)#$$*[ %# ] Ketsch, Hohwiesen(D) Openairriverterrace Weichselian (glacial) "!?unclear[ %# ] Kleinbesten(D)Openairriverterrace Weichselian (glacial) "#?unclear[ %# ] K onigsaue(D) Openairriverterrace (hyenaden/overlapping Neanderthalcampsite) Weichselian (glacial) #&(skull)!?unclear[ '" %# ] K orbisdorf(D)Openairriverterrace Weichselian (glacial) "$?unclear[*/ K orbiskrug(D)Openairriverterrace Weichselian (glacial) "#?unclear[ %# ] Kon eprusyCave (CZ) Carboniferouslimestone cave(hyenapreydepot den) Weichselian (glacial) #!( !' coprolites,##$ bonesfrom individual skeleton) !&%##[ &) ] KreuzCave(D) Carboniferouslimestone cave(unclearsite,?cave bearden) ?Weichselian (glacial) "!?unclear[ %' ] LahntalCave(D)Karstcave(cubraisingden) Weichselian (glacial) )!"#&[ %' ] Lampertheim,In derTanne(D) Openairriverterrace Weichselian (glacial) "!?unclear[ %# ]

PAGE 18

#) PaleontologyJournal T7894#:Continued. LocalitySitetypeAge Hyenas (NISP) Lions (NISP) Bone assemblage (NISP) References Lampertheim, L uderitzbucht (D) Openairriverterrace Weichselian (glacial) "#?unclear[ %# ] Lichtenau, Hasenkopf(D) Openairriverterrace Weichselian (glacial) "#?unclear[ %# ] Lipperode(D)Openairriverterrace Weichselian (glacial) #"?unclear[ $* ] MartinsCave(D) Carboniferouslimestone cave(cavebearden,hyena cubraisingandcommunal den,wolfden) Weichselian (glacial) #$*#* #%(realamount unclear),many cavebear remains [ %' ] MedvediaCave nearLiptovsk y (Sk) Limestonecave(cavebear den) Weichselian (glacial) Skeletonofa subadult ?unclear[ )% ] Minice(CZ)? Weichselian (glacial) "#?unclear[ )" ] M ucheln(D)Lake ?Eemian/ Weichselian "#?unclear[ %# ] Neumark-Nord Lake#(D) Lake,forestelephantsata graveyardsurroundedby MiddlePalaeolithiccamps Eemian (interglacial) % ($ coprolites) $ (and#*!bones fromilllioness individual skeleton) Many hyena/lion scavenged elephant carcasses [ '% )$ ] Niederlehme(D)Openairriverterrace Weichselian (glacial) #*?unclear[ %# ] OberneissenCave (D) Karstcave(cubraisingden) Weichselian (glacial) #$"?unclearunpublished Oberr oblingen (D) Openairloess Weichselian (glacial) "#?unclear[ %# ] Oberrohn(D)Openairloess Weichselian (glacial) "#?unclear[ %# ] Otterstadt, Altrhein-S ud(D) Openairriverterrace Weichselian (glacial) "!?unclear[ %# ] Otterstadt, Altrhein-Nord (D) Openairriverterrace Weichselian (glacial) "&?unclear[ %# ] Otterstadt, Waldwiesen(D) Openairriverterrace Weichselian (glacial) "&?unclear[ %# ] Osteroden(D) Zechsteingypsumkarst hyenaden(typeunclear) ?Weichselian (glacial) #"?unclear[ '" ] PerickCaves(D) Carboniferouslimestone cave(hyenacubraisingand communalden) Weichselian (glacial) #$*('skulls)'/&*&[ #" %! ] Petershagen(D)Openairriverterrace Weichselian (glacial) "# Woolly rhinoceros skeleton, unscavanged [ %# )) ] Pfe+erburgCave (D) Carboniferouslimestone cave(hyenaden,cavebear den,?temporarywolfden) Weichselian (glacial) &*(and) coprolites) &#(realamount ?unclear) [ %' ] Phoeben(D)Openairriverterrace Weichselian (glacial) "#?unclear[ %# ]

PAGE 19

PaleontologyJournal #/ T7894#:Continued. LocalitySitetypeAge Hyenas (NISP) Lions (NISP) Bone assemblage (NISP) References Praha-Podbaba (CZ) Openairriverterrace Weichselian (glacial) #%(fromtwo individual skeletons) ?unclear[ )" ] Praha-Ko s re (CZ) Openairriverterrace Weichselian (glacial) "#?unclear[ )" ] Praha-Libe n(CZ)Openairriverterrace Weichselian (glacial) "%?unclear[ )" ] Questenberg(D)?unclear ?Weichselian (glacial) #"?unclear[ '" ] R osenbeckCave (D) Carboniferouslimestone cave(hyenacommunaland cavebearden) Weichselian (glacial) )&('skulls)" ?unclear,many cavebear remains [ #* ] Selm-Ternsche (D) Openairriverterrace Weichselian (glacial) &"?unclear[ $* ] Siegsdorf(D) Riverbranch,riverterrace site Weichselian (glacial) ( &coprolites) %!bonesofone maleindividual skeleton includingskull Mammothbull carcass scavengingsite [ *' ] Speyer,Deutschof (D) Openairriverterrace Weichselian (glacial) "#?unclear[ %# ] Svobodn eDvory nearHradec Kr alov e(CZ) ?openair Weichselian (glacial) "#?unclear[ )" ] Sewecken-Berge Quedlinburg(D) Triassicgypsumkarst hyenaden (preydepotandcommunal den) EemianWeichselian (interglacial, mainlyglacial) ##'( #skullof cub) #*%%"[ '! %# ] Senzig(D)Openairriverterrace ?Weichselian (glacial) "#?unclear[ %# ] SloupCave(CZ) Carboniferouslimestone cave(hyenaden,cavebear den,?wolfden) Weichselian (glacial) )'(!skulls)$'![ $" ] Sophie'sCave(D) Jurassiclimestonecave (hyenacubraisingand communal,cavebar,and wolfden) Weichselian (glacial) #! % ,manycave bearremains (o.encarnivore damaged) [ $' ] Speyer, Binsfeld-S udost (D) Openairriverterrace Weichselian (glacial) "#?unclear[ %# ] Srbsko Chlum-Kom n Cave(CZ) Carboniferouslimestone cave(hyenapreydepotand commutingden) Weichselian (glacial) &%%(from% individual skeletons, including cubs,$ coprolites) Twoindividual skeletons:#$/ bonesfroman adolescentill lioness,and#"* bonesofa juvenile /$*(mainly Przewalski horseremains) [ &/ ] Teufelskammer Cave(D) Carboniferouslimestone cave(hyenaden,cavebear den,?temporarywolfden) Weichselian (glacial) *##/*[ $# ] ( iede(D) Zechsteingypsumkarst hyenaden (preydepotandcommunal den) Weichselian (glacial) &"(#skullof adult) #$ #!"(evenmore, unclearamount atthisstage) [ %* ]

PAGE 20

!" PaleontologyJournal T7894#:Continued. LocalitySitetypeAge Hyenas (NISP) Lions (NISP) Bone assemblage (NISP) References T urmitz(CZ)Openairloess Weichselian (glacial) !(#skull)!?unclear[ #* ] # Ust nadLabem (CZ) Openairriverterrace Weichselian (glacial) "#?unclear[ )" ] V ypustekCave (CZ) Carboniferouslimestone cave(hyenaden,cavebear den,?temporarywolfden) ?Eemian/ Weichselian (interglacial/ glacial) !&%(&skulls, one individual skeleton) &%?unclear Unpublished, skeletonin[ &/ ] Wanne(D)Openairriverterrace Weichselian (glacial) "#?unclear[ %# ] Westeregeln(D) Zechsteingypsumkarst hyenaden (preydepotandcommunal den)andoverlapping Neanderthalcampsite Weichselian (glacial) )$( #"skulls, and!" coprolites) #!$))[ '# %# ] WildkirchliCave (Ch) Limestonecave(cavebear den) Weichselian (glacial) "!& ?,manycave bearremains unpublished Wiesental, Allmendweg(D) Openairriverterrace Weichselian (glacial) "#?unclear[ %# ] Wiesental, Viehweg(D) Openairriverterrace Weichselian (glacial) "#?unclear[ %# ] WilhelmsCave (D) Carboniferouslimestone cave(hyenacubraisingand communalden) Weichselian (glacial) #%/ #'(sibling skeleton remain) #!%[ %' ] W orth,Geisb ogel (D) Openairriverterrace Weichselian (glacial) "#?unclear[ %# ] W orth, Rheinanlagen(D) Openairriverterrace Weichselian (glacial) "#?unclear[ %# ] W usteScheuer Abri(D) Zechsteinlimestone abri/smallcavity Weichselian (glacial) !*( #skull)"%#[ '$ ] Zechovicenear Volyn e(CZ) ?openairLoess Weichselian (glacial) "#?unclear[ )" ] ZoolithenCave (D) Jurassiclimestonecave (hyenacubraisingand communal,cavebar,and wolfden) Saalianto Eemian/ Weichselian (interglacial/ glacial) !"*(!skulls)!!/(/skulls) Fewbones, mainlydamaged cavebearbones inunclear amount(about halfMillioncave bearbones) [ $& $$ ] ( .OntogenyandSexualDimorphism. ( eontogenyin C.c.spelaea wasbestknownonlyfromitsteeth,which weredeterminedwithmilk,cub,andadultteethofalljaw positionsbyReynolds#/""[ #/ ].Here,theontogeneticstages aredemonstratedfortheskullwiththreestages,(a)sibling withmilkdentition,(b)cubwithfullychangedpermanent teeth,(c)andadultwithalreadywornteeth( Figure! ).Also compiledareskeletonsinsibling(compositeskeleton)and adultages(individualskeletons, Figure# (d))fromCzech Republiccaves.Fromsomecavesmilkteethanddentition suchasafewrarebraincasesandnearlycompleteskullsare gured[ #/ !$ !% !/ &! &* &/ $# $' '# ].O.enpostcranial siblingandcubremainsweremisidentiedinoverlapping hyena/cavebeardens,becausetheirboneslookverysimilar, andalsothebraincases.Postcranialboneshavebeen-gured fromafewcavesandopenairsitesofEurope[ $# $' '# ], becauseo.enthosearestronglydamagedandremained o .enunrecognizedwithfragmentsinthepreybonemartial. ( esexualdimorphisminmodern[ /$ /* ]andPleistocene hyenas[ #* )/ /# ]ingeneraliswellknownwiththelarger females,whosestatisticsarepresentedforseveralcavesof GermanyandCzechRepublic[ $' ],whereasinmanycases thesexidenti-cationremainsunclear,especiallyifthesite isnotwell-datedtobeinterglacial(smallerforms)orglacial (largerforms),orinthecloseoverlapofsmallfemalesand largermales.

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PaleontologyJournal !# % .Cannibalism. ( edocumentedandillustratedrecord ofbonedamageintheLatePleistocenespottedhyenasis recentlymuchbetter[ '/ ]thanthemodernrecord.(ere aretwotypesofbonedamage:(a)crackedbones,which even-t,aredemonstratedbestfortheSrbskoChlum-Kom n Cave(CZ)[ $# ]whereanoldhyenacarcassinthedencave itselfwasconsumedandcrushedintopieces( Figure& (b));(b) chewedandnibbledboneswithmanybitemarks,especially atlongbones,andcrackedlowerjawsaremoretypicalat communalandbirthdensites[ &/ $& '# ].Atcavesites, postcranialhyenaremainsaremorecommon,whereascranialremains(especiallyskulls)aremorerepresentedatopen airsites(e.g.,BadWildungen,Bottrop,UpperRhineValley sites,Westeregeln,andothers)[ $) '% ],whichissimilarto somemodernspottedhyenaopenairdensinAfrica[ /$ /% ].Hyenasdenscontainsimilarlydamagedbones(similar damagesastheirpreybones).(eremovalofthelowerjaws oftheirownspeciesskullswasonlypossibleduetobreakage ofthejugalarchesandbreakageofthejawsbehindthelast teeth;thereforelowerjawsareveryo.enfoundwithoutthe ramusandskullswithoutjugalarches[ #* '/ /* /) ],which issimilartodamageonlionorcavebearskulls.Scavenging isreportedinallageclassesfromseveralcavedens,whereas eveninfantandcubremains(cranialandpostcranial)very o .enshowpatternsofcannibalismdamage,similartothat onjuvenileremainsfromamodernAfricanspottedhyena den[ &/ ].(ecommonkillsespeciallyinsiblings[ // ]are a .ernewstudiesofmodernspottedhyenasnotrelatedto thematernalrankandnotcorrelatedwithcubsurvivalinthe modernspottedhyena[ #"" ].( edamageexhibitedoftheir ownspecieslongbonesissimilartoothercarnivores(lions, wolves)andalsotocavebears[ %# ]. '.".DenTypes. ( erearethreedi+erentLatePleistocene typesofdensinEuropetodistinguishbasedon(A)landscape morphology(Figures and ) ):(a)cavedens,(b)gypsum/limestonekarstopenairdens,and(c)riverterrace/loess openairdens[ &/ '# '! '$ '% ]whicharemappedin detailinthecasestudyareaofGermany/CzechRepublicin both,lowlandsandmountainousregions(Figures # and ) ). Hyenapopulationstructureandboneassemblageanalyses (botha.erNISP)suggestthatthosemorphotypesofdens arecomparabletomodernAfricanspottedhyenadentypes similarlybasedonthe(b)ethologyanddemographyof populations[ #"# #"$ ],whichismoredi: culttocompareto palaeopopulationsandtheirfossilrecord.( reemaintypes aredistinguishedintheLatePleistoceneofEurope( Figure) ): (a)birth/natalden(similartoAfrica[ #"& ]:siblingbones/milk teeth,abundant"nibblingsticks",andfewpreyfaunaremains whicharestronglydamaged,e.g.,[ &* $& $/ '& ]);(b) communalden(similartoAfricaandmostcommondentype [ !$ &' &/ $# $& $' '& '$ ]:cubandmainlyadulttosenile hyenabones,o.encannibalisticdamaged,preyremainsmore fragmentedandchewed,andabundantcoprolitesforden marking);(c)preydepotden(mainly/onlyadultremains andevenarticulatedskeletons,abundantpreyremainspartly bodypartsinarticulationandlessbonedamage,andfew coprolites),whichismoretypicaltypeoftheLatePleistocene coldperiodandcave-richregionsofEurope[ &$ $" ]. $.PreyRemain"Bone"Accumulators. Hyenidsofthe MiocenetoEarlyPleistocene(includingdi+eentecomorphs: bonecrushing,hunting,civet-like,andothers)seemtohave startedtodevelopboneaccumulations(especially Crocuta ) withbonedamageduetopreyimporttodens(andless singlebones).(eearliestEuropean Crocuta recordsdate backtothePliocene[ #"! #"& #"' ]withfewsitesknown fromtheEarly[ #"% #") ]andMiddlePleistocene[ #"/ ].Only bone-crackingecomorphhyenidspersistedinEuropetothe LatePleistocene.(emostfamousEuropeanEarlyPleistoceneUntermassfeld(D)sitewitha Pachycrocuta population [ #") ]hereidentiedasaverytypical"hyenaopenairand boneaccumulationdensite"onariverterraceposition(high amountof Pachycrocuta remains,coprolites,manychewdamagedpreybones,o.enlegboneoverrepresentation),with possiblelocalshort-distancetransporta.erthecarnivore preyremaindepositionswasevenincorrectlyinterpreted asresulting"onlyby, oodsaccumulated"boneassemblage, whichdemonstratethatopenaircommunalhyenadensites arestillo.enoverlooked.Ashereincompiled,thebest andmostdensefossilrecordoflargecarnivoresandtheir activitiesandboneassemblagesisfromtheLatePleistocene ( Figure) ).Tounderstandthosefossil"carnivoreboneassemblages"mostbonetaphonomystudiesdealtmorerecentonly withthequestionabout"anthropogenicorcarnivorebone assemblageidentication"[ ##" ##% ]butdidnotconsiderthe palaeoecologyofextincthyenasandthecomplexcavebone taphonomy,especiallyincaves.Boneaccumulationscaused bymodernspottedhyenas,toavoidcon,ictwithlionsand othercarnivoresandtofeedtheircubsatthedensite,are studiedinseveralsmallcavesandopenairlocationsinAfrica insomecasesatleastbytheirNISPandMNI[ /% #"* #!! ]. InEuropetherewasnotastandardtoanalysehyenadenprey boneassemblages,ando.enonlysingleanimalgroupswere publishedfromsitesandarelistedwithoutNISPandMNI reports[ &" ],whilemorerecentworksincludethepreybone quantitativeanalyses.(esequantitativestatisticsallowa-rst generalhyenadensite(versushumansite)identi-cation, because,atmostLatePleistocenespottedhyenadensites, #"&'%ofthebonesarehyenaremains[ &! '$ ]( Figure) ). Inasecondqualitative/quantitativestudy,lessthe"classical bitemarkanalyses"[ #!& #!$ ]supportsahyenadenbone accumulationorigin,asfurthermorethe"repeatingbone damagestageanalyses"[ #!' #&" ],whichisbestmadewith elephantandrhinocerosremains[ '" '* ]andboneelement abundance[ #!% #!* ]whereastheschleppe+ect[ #!/ ](= selectionofbodypartsatscavengingsiteandnonimportation ofalltypesfrompreybody,suchasvertebralcolumns)is includedinsuchstudies.Finallythegeneral"damagedegree" isimportantsimplytodistinguishhuman"kitchenrubbish" (=mainlybonefragmentsatsites),and"carnivoresites"(= abundantcompleteandpartlyarticulatedremains)[ ##" #&" ].

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!! PaleontologyJournal '.).BoneDamageStages. ModernandLatePleistocene spottedhyenasare"bonecrushers"[ !! #&" ]withtheirspecializedbonecrackingjawsanddentition[ )/ #&# ]( Figure% ). Remainsofoldagedindividualsshowcrowndamageorthe lossofmainlythelowerjawpremolarteeth(bonecrushing teeth)andfullyusedincisiveteeth(bonenibblingteeth) whichisdemonstratedbothformodern[ #&! ]andnowfor theLatePleistocenespottedhyenasofEurope.(ereare twomaingroupsofbones/preytypes:(a)elephantsand rhinoceroswithunbreakablebonesand(b)bovid,equid, cervid,ursidandantelopeherbivorous,andcarnivorebones whichcanbecrushedbyhyenas.Forextinctelephant( Mammuthus [ '* ], Palaeoloxodon [ ') ])andrhinoceros( Coelodonta [ '" ]),ineachtaxon,threebonedamagestageswerepresented beingnearlyidenticalasthoseofsimilarhyenabutchering/decompositiontechniquesonthelargestpreyandof similarspongious-lledbonestructureof"uncrushablemassivebones."InotherLatePleistocenemegafaunamammal prey,thosestagesarenotyetwelldistinguished,suchas forhorses,bovids,orcervids,buttypicallymoreabundant distallegelementswerele.o.enuntouchedatdensites[ '& '' #&& #&$ ],similartothoseknownfrommodernspotted hyenadens[ #!' #!% #!/ ].Atsomesiteseventhosedistalleg bones(metapodialsandphalangesmainly)ofdeers,horses, orsteppebisonswerealsocrushedasbestdocumentedat thePerickCaves(D)hyenaden[ #&$ #&% ]orSrbskoChlum Kom nCave(CZ)[ $# ]andFuchslukenCavity(D)[ #&* ].Only twostagesaredescribedformodernbonesha. sandbone fragmentsofcrackedlongbones.Alsothecervidboneswere muchmoreeasiertocrushintopieces(similarasantelope bones[ #&% ]),and,therefore,longbonesaremostlypresent fragmentedandnotchewedatLatePleistocenespottedhyena densitesalloverEurope.(enonelephant/rhinocerosbones were-nallyo.enusedas"nibblingsticks"[ #&" #&% ]. '.!.NibblingSticksandPlayBones. ( esocalled"nibbling sticks"areverytypicalatbirthdensandlesscommonat communaldens,thoughabsentatpreystoragedens[ &/ ]. (eseareanykindofbonefragments(mainlynonelephant/rhinocerosbones)whichwere-rstunipolar,then bipolarchewedmainlybysiblingsandcubsforteething purposes.(eyarebestrepresentedattheNadKa cakem Cave(CZ)[ &/ ]wheremorethan%"stronglychewedbone fragmentshavebeenfound.(esenibblingsticksinclude lessmammoth[ '* ]andrhinoceros[ '" ]andareknownfrom otherLatePleistocenespottedhyenacaveandopenairden sitesasusefulforhyenadenidenticationbutarenotyet publishedwellfrommodernsites,however.Besidesthose, playbonesarepresent,especiallyofelephants[ '* ') ],which arepiecesmainlyfromthepelvisandscapula(="Neanderthal pseudohandaxes"[ '* ])orfemurjointheadsofyoung elephants[ ') ]whichwerealsostronglychewedoverlonger periods. &* .DenMarkingbyFaeces. ( efossilrecordisproblematic, becausecoproliteshavenotbeenrecoveredorcrumbledto "dust"overthecenturiesinmanycollections.Hyenadensand territorialboundariesaremarkedbyextanthyenas[ #&) ]and weremarkedintheLatePleistoceneasdocumentedbestat communaldensites[ %" #&/ ].InsomeEuropeancaveshyenas trampledevencompletephosphatichorizons,historically describedforLindentalCave[ #$ ]orK onig-LudwigsCave (D)[ / ],whereasinSrbskoChlum-Kom nCave(CZ)the phosphatepiecesbuiltahighpercentageofthesediment[ $# ]. AtCzechSloupCave,SrbskoChlum-Kom norKon eprusy Cave[ $" $! ],andFrenchcaves(e.g.,RochelotCave[ &$ ] andothers)orSpanishcaves[ #$" ],coproliteshavebeen reportedtobepartlyabundant,whereasalsoattheopen airsiteWesteregeln(D)twofaecalplaceswithcoprolite concentrationsweredocumentedatanopenaircommunal gypsumkarstdenwithinthesinkhole[ %" ].Anoverviewof thefewsurvivingmaterialhasbeenpublishedfortheGerman andCzechhyenadenandcarcassscavengingsiteswhereas thedi+erenttypicalhyenashapedpelletsbuiltaggregates ( Figure& (c)),whicharewellcomparedtomodernspotted hyenaexcrements[ %" ].Modernhyenasmarktheirdensand territoryagainstotherclansandevenlions[ #&) #&/ ]which mustbeexpectedforthePleistocenerecord,too.Suchearly cementedcoprolitescansurviveevenwatertransport[ #$# ], especiallyifcementedandencrusted,forexample,bycaliche, andevensurvivesaltwaterasreportedfromtheNorthSea [ #$! ].Extractionsofpollenfromsuchexcrementswereused toreconstructthevegetationandpalaeoenvironment[ #$& ], butthosestudiesdidnottakeintoaccountthattheseplant remainsarenotconsumedbyhyenasandseemtoresult morefromintestine/innerorganfeedingbyhyenasoftheir prey.(erefore,thepollenfromhyenafaecescancompletely re,ectthedietofhyena'sprey(=lastplantfoodoftheprey), whichitselfwasselectiveinplantfeeding[ %" ].Toreconstruct "vegetationandlandscape"basedonlybythisisproblematic, butincombinationwithhyenapreyanalyses,itmighthelpto understandthepreyofhyenasusingtheircoprolitesbasedon pollenandbonefragmentDNAanalyses. && .HorseandDonkeyHunters. ( emainpreyfoundat hyenadensitesalloverEuropeisfromhorses.According tothelatestrevisionoftheLatePleistoceneEuropeanhorse Equusgermanicus Nehring,#))$"basedonthosefrom Westeregeln(D)hyenaden,itisdeterminedthattheyare synonymoustothemodernsmallPrzewalskihorse Equus caballusprzewalskii Poljakov,#))#[ '& ].EvenattheSeweckenBerge(D)hyenaden,the"unicornholotypeskeleton"was demonstratedtohavebeencomposedofPrzewalskihorse skeletonremains[ '$ ].Atthissitehyenashuntedadditionally a"largerhorse" Equuscaballus cf. fossilis (taxonomystill unrevised)withintheEemianinterglacialtimes[ '$ ].At severalcaves,includingtheWookeyHolehyenaden(GB) [ #& ],horsesarethemostabundantprey.Atdensitesabundantteethwerefoundwhichresultedfromthecrushingof thethin-walledskullsandmoderatelymassivelowerjaws. Additionally,alsoasbestdocumentedatRochelotCave[ &$ ], bonesatdensitesaredominatedbydistallegelements,which wherefoundpartlyinanatomicalconnections(=legimport). (ehorsehuntingspecializationoftheLatePleistocene spottedhyenasisbestandmostimpressivelydocumented attheSrbsko-Chlum-Kom nCave(CZ),where'# %ofthe

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PaleontologyJournal !& NISParesmallcaballoid E.c.przewalskii horsebones(more than#,'""bones).Formodernspottedhyenas,equid(zebra) predationiswelldocumentedattwoAfricanhyenadenbone assemblagesoftheAmboseli[ ##/ ]andSyokimanu[ /% ]dens sites(!&%and#&%zebras)andzebrascanreachupto*"% ofhuntedremains[ #&& ].WithintheSrbsko-Chlum-Kom n Cave,remainsofthreefoetalhorses(onenearlycomplete foetusskeleton)suggestedthathuntingtimeoccurredin thespringtime;thereforemotherhorseswereimportedas carcassestotheden[ #&& ].(eLatePleistocenerecordthere indicatesthatthehuntwastargetingmainlygrownuphorses ( /!%),whichissimilartozebrahuntinginAfricanhyenas [ #&& ].( ehuntingofhorsesdoesdependonthedi+erent LatePleistocenelandscapes( Figure) (b)),astheyaremore abundantinmammothsteppeenvironmentsbutcanbe highlyabundantinvalleysofmountainousregionswhere therearemuchfewercavebearsasapossiblefood(e.g., BohemianKarst[ &/ ]).(isagainissimilartomodernhyenas, whichalsodependonlandscapedi+erences(woodland, mountain,andsavannah)fordi+erentprey,andtherefore preyabundancecanbeverydi+erentinmodernspotted hyenaboneaccumulations[ '/ )) /' /* #"% ##' ##* #!! #&& #$$ ].LatePleistocenedonkeys Equushemionushemionus werealsotargetedmainlyinthesteppeenvironments,as demonstratedwellbytwositeswhichhavetheirremainsin thehyenadens,theFuchslukenCavity(D)[ #&* ],andAgios GeorgiosCave(Gr)[ !/ ].Inothercavesandopenairdensof Europe,thesedonkeysaregenerallyrarerinthehyenaden boneassemblages(#&%). &# .BovidHuntersorScavengers. ( esteppebison Bison priscus wastargetedinthecoldperiodsonlyinsomeareasor periodsbyLatePleistocenespottedhyenas,especiallywhere hyenadenswereabundant,butcavebears(andelephants) wereraresuchasinthe(uringianKarst[ '% #&* ].( ereare onlyfewexampleswheresteppebisonhuntingspecialization developedasreactiontocavebear/mammothpreyabsence orrareness,suchasatFuchslukenCavity(D)[ #&* ].Atthis hyenaden,ahighabundance(&!%NISP)ofbovid,mainly steppebison,isuniquefortheGermanandCzechhyena denrecord.InCamiacCave(F),thesteppebison(and/or bovid)remainswerecalculatedtorepresent&/%oftheprey fauna[ !* ].AlsoinRochelotCave(F),bovidswerethesecond abundantpreyremain[ &$ ].InItalySanTeredoCave(Sicily) evenhigheramounts(upto'"%NISPinsomelayers) werefrombovids[ &* ],alsoindicatingaspecializationthere. Generally,steppebisonsarerepresentedinsimilarabundance ashorsesinthemammothsteppeboneassemblagesand aremorerareinborealormountainforesthyenadenbone assemblages( Figure' (b)).(elessimportantrole(fewNISP) ofsteppebisonatmosthyenadensitesinlowlanddensites (e.g.,Westeregeln,Bottrop)alsoindicatesacompetition/prey overlap,probablywiththeLatePleistocenesteppelionsand wolves,whichweretheactivehunters,hyenasmorethe scavengers( Figure) (b)).( issuggestionisalsosupported byrarebisonNISPamountattheSrbskoChlum-Kom n Cavehyenapreydepot,whereasinthismostlargeEuropean hyenadenboneassemblage(NISP=&,%/')onlyfewsteppe bisonremains(!%ofNISP)arefound[ $# ].Also,inthesame landscapeatthehyenapreydepotofKon eprusyCave,only % %oftheboneassemblagematerialwasfromthesteppe bison[ $" ].(ehabitatvariabilityinmountainregionswith limitedcavebear/mammothseemstohavedeterminedeither thebovidorequidhuntingspecializationandlogically thosehadtobehuntedinhigheramountstoreachthe nutritionbiomass.(isexplainsthebesthighbovid/equiid preypercentagesatbothsites(FuchslukenCavityandSrbsko Chlum-Kom nCave).( isissimilartoAfricanlions,which alsospecializeinhuntingofbu+aloinwetlandareas(e.g., AfricanAmboselihyenaden)[ ##/ ],whereasCanadianwolf clanstargetwoodlandbisoninmountainwoodlandregions ofCanada[ #&* #$' ].( esamehyena/lion/wolfpreyrole occurs,thoughmuchlessfrequently,with Bosprimigenius mainlyinwarmageperiods.Scavengingiscomparableto themodernspottedhyenasscavengingin"elephant-free" areasasdocumentedintwomodernAfricanhyenaden boneassemblagesoftheAmboseli[ ##/ ](bu+alo/domestic cattle=% %,wildbeest=&&%)andSyokimanu[ /% ]denssites (domesticcattle=&"%,wildbeest=* %)[ #&& ]. '.&'.DeerScavengesandShedAntlerCollectors. Fastrunning cervidswerenotamainhuntedfoodsourceforslowspotted hyenas(Figures ) (b)and ) (c))similartothatgazellesarenot themainhunted(butscavenged)preyinmodernAfrican spottedhyenas[ '/ )) /' /* #"% ##' ##* #!! #&& #$$ ]. ( eirrarityinthebonerecordofallstudiedhyenaden boneassemblagesmustbeinterpretedcarefully,becausetheir bonesareeasytocrushandtoswallowforbonecollagenuse. ForthePerickCaves,the Megalocerosgiganteus bonerecord demonstratesthatcranialremainsanddistallegelements dominatethebonerecordatdensites[ #&% ],asrepresentedat theSewecken-Berge[ '$ ]andFuchslukenCavity[ #&* ].Atall sitesstudied,therecordofredandgiantdeerisalwayslimited andbonesarehighlyfragmented,whereastheremainsof reindeerarevariableinamountsandpreservation,especially athyenadens/overlappinghumancampsites[ '& ].Compared tobovidsandequiidsorrhinoceroses,isolatedteethofthe huntedcervidpreyaremoreusefultoestimatetheMNI.Even withthis,cervidsarerarelyrepresentedbythetoothrecord atthedens.Obviously,cervids( Megaloceros,Cervus,Dama, and Capreolus )werenotanimportantormainfoodsource fortheLatePleistocenespottedhyenasatall(e.g.,nearly absenceeveninborealforestdensiteslikeSrbskoChlumKom nCave[ $# ],wherereindeerremainswerepossibly importedbywolves).Veryinterestingandonlytypicalfor hyenasintheEuropeanLatePleistoceneisthecollectingof sheddeerantlers,whichispartlycomparabletocollected hornsofgazellesfoundatAfricandensites[ /% ].( estudy ofallhyenadenantlerrecordsfromLatePleistocenedens incentralEuropeindicatesthattheimportationofantlers (##"antlersonlyoneachdensite)byhyenasalwaysle. similardamagemarkstotheremains:thebasewithstrong bitescratchmarksatthechewedendandthoseonantler fragmentsofolderindividuals.(eroughlowerattachment partwasneverchewed,becauseitcouldhavedamagedthe toothmeat.

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!$ PaleontologyJournal '.&(.WoollyRhinocerosCarcassDecomposers. Onlyasingle woollyrhinoceroscarcasshasbeenfoundinPetershagenof northernGermanyandwasfoundinarticulationuntouched bycarnivores[ #$% ].(ereisonescavengedskeletonfrom theKr olpa(D)openairgypsumkarstscavengingsite[ '% ] andmostprobablyanotherskeletonfromtheBadWildungen hyenaopenairloessden[ '# ].Bothcarcassesdemonstrate carcassdecompositionstage!(of&),wheretheanatomical contextismoderatelytomostlydestroyed,andpartshave beenalreadyremoved.Inmosthyenadenssitesthebones areisolatedorbodypartsarepartlyinanatomicalcontext (mainlylegs).(edamageonrhinocerosbonesoftheLate Pleistoceneisthebest"hyenadenmarker"atall,because bonesfoundatdensoropenairscavengingsitesfollowa consistentdamagepattern[ '" ]. Coelodontaantiquitatis skulls (e.g.,Kon eprusyCave,SrbskoClum-KominCave,Kr olpa) [ &/ $# '% ]aredocumentedtohavebeendamagedsimilar ascannibalisticdamagedhyenaskulls(crackedjugalsand ramusdamage).Insituationssuchascommunaldensites, theskullsexhibitevengreaterdamage,andonlyteethare le ,andinseveralcaseswereevencompiledindentition rows(e.g.,Westeregeln[ '& ],Sewecken-Berge[ '$ ]).(ere andatopenaircarcassscavengingsites,hyenasle.excellent examplesofbraincaseopeningoncalftoadultindividual skulls(BadWildungen[ '# ],Selm-Ternsche[ '" ],andKr olpa [ '% ]).AlloverEurope,LatePleistocenespottedhyenascaused threemainandsimilarbonedamagestagesonthepelvis, scapula,andlongbones.Sucharedocumentedingreater amountsatopenairdens(Bottrop[ '" ],BadWildungen[ '# ]) orcavedensites(LindentalCave[ #$ ],SveduvSt ulCave[ !$ ], TeufelskammerCave[ $& ],BalveCave[ $$ ],SrbskoChlumKom nCave[ $# ],Kon eprusyCave[ $" ],orSloupCave[ $! ]). Withthecarcassscavengingsites(carcassdecomposition stages#&),abutchertechniqueforcarcassdecompositions onthesecondlargesthyenapreyisnowdemonstrated,which onlyishyena-related.(ishasnotbeenstudiedyeton modernAfricanrhinoceroscarcasses.Withintherhinoceros carcassbutcher/decomposingtechnique,hyenasmovedbody partstothedensitestoavoidcon,ictswithlions/wolves. ( isisnearlyidenticaltothecarcassdecompositionand bonedamagestagesofelephantbones(mammoth,forest elephants).Atmanycavesites,thewoollyrhinoceroswas importedabundantlywhichindicatethatthesewereimportantfoodsourcesformeatandbones(e.g.,Teufelskammer Cave[ $& ],Bottrop[ '" ],BadWildungen[ '# ],Westeregeln [ '& ])beingmostlyonsecondpositionwithintheNISP(e.g., LindenthalCave[ #$ ],WookeyHole[ #' ],andSewecken-Berge [ '$ ])(Figures / (b)and / (c)),alsoasresultofrobustnessofthe uncrushablebones. &% .WoollyMammothCarcassButcherTechnique. ( e decompositionofextinctandextantelephants( Loxodonta [ $% ], Palaeoloxodon [ ') ],and Mammuthus [ '* ])bymodern andLatePleistocenespottedhyenasisnowbeingstudied indetail.Lionsandhyenasfeedingonelephantcarcasses overlap( Figure/ (c))butarerecognizedonlyonceinthe initialstage#ofcarcassdecompositionofawoollymammoth attheSiegsdorf(D)bullcarcass(intestine/innerorgan,trunk, andfeetfeeding[ '* ') #$* ]).(edestructionofthebones issimilartorhinoceros,becauseofsimilarpreysizeand similarbonestructure.Inelephantandrhinoceros,thebones are-lledbyspongiosaandarenearlyunbreakable,even forhyenas[ '* ') ].(eyonlycanchewtheirlongbones andlargerbonesstartingfromtheso.distaljoins.(e elephantfeedingstartssimilarlyinbothtoppredatorsquite o .enontheinnerorgans/intestines,whichbodycavitythey reacho.enovertheanus[ #$) ].Caninebitescratcheson theinnersideofthethoracic/lumbarvertebralcolumns onthree Palaeoloxodon skeletonsofNeumark-NordLake# demonstratedthattoppredatorswentintothebodycavity eatingfrominside[ ') ],whichisevenknownfrommodern hyenasandlionsindocumentary-lms[ #$/ ].(isissimple toexplain,becausetheverythickskinisevenhardtocut bytoppredators.(eskinattheanusisthinandfeeding frombehindallowstoconsumeintestines-rst.Alsothetrunk isinitiallyeateninelephantcarcassesorthefeetmainlyby hyenaswhichwasmonitoredatanAfricanelephantcarcass [ #$) ].( erearethreemaindecompositionstagesonelephant carcassdocumentedinthefossilrecord.Hyenasaretheonly predatorsthatstartthesecondstageonelephants,especially onthelegs.Withthis,thecarcassisdamagedanddemonstrateswhythereareonlyfewmammoth/forestselephant skeletonremainsfoundcompleteinEurope.Exceptionsare carcasseswhichwerefoundinshallowlakesorriverbranches partlycoveredbywater,suchasatNeumark-NordLake# (severalskeletonsof P.antiquitatis [ ') ]).(ebestexample ofinitialfeedingon M.primigenius isthelargebullfrom Siegsdorf(D)[ '* ].( eAhlen(D)orKlinge(D)skeletons arefurtherdamaged(decompositionstage![ '* ]),whereas inallthereisalsotypicaldamagetotheskull,whichis thin-walledandspongae-likeinconstructionandeasyto damagebyhyenas.(e-naldamageofthelargerbones happenedatcommunalorbirthdensiteswheremainlybones ofmammothcalvestosubadultswerefound[ '* ],simply areasonoftransportationselectionofsmalleranimaland bodypartsoflessweight(=schleppe+ect[ #!/ ]).Insome caseshyenasfedonmammothremains,whichNeanderthal humansle.atcavecampsites(WeinbergCaves[ #$/ ]),or stolefragmentswhichtheyimportedtotheirdens(Perick Caves[ '* ]).(egreateramountofcalfremainscannotonly besimplyinterpretedas"huntedmammothcalves"byhyena clans,whichindeedcannotbeexcluded.Modernlions(or evenLatePleistocenesubspecies)arethemoresuccessful elephantcalfkillers[ #$) ].Inallboneaccumulationsathyena densmainlyteethandbonesfrommammothcalvesand subadultshavebeenfound,whereastheirpercentageinthe NISPisalwaysnotthathigh( Figure/ (b)),butthebodymass andweightareindeedmuchhigher.Obviously,elephants werelowlandinhabitants,andthereforetheyareveryrare orabsentinborealforesthyenadencavefaunalassemblages (BohemianKarst[ &/ ],(uringianKarst[ #&* ],andSauerland Karst[ '* ]),becauseofseasonalmigrationwithinthevalleys ofthemountainslopes. '.&".CaveBearScavengersinCaves. ( escavengingofcave bears(andbonedamage)wasinitiallybelievedtobea resultof"cannibalismwithincavebears"[ ## '! ].( iscan onlybeunderstoodbyincludingmegafaunaquantitativeand

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PaleontologyJournal !' qualitativeanalysesandthehyenaandlion(alsowolf)bone record,includingtheirtaphonomyandethology[ %# ].Many argumentsagainstthis"bearcannibalism"hypothesishave beenpublished,suchasthetoothmorphologychangesas adaptationstoplantfeeding(increasingofenamelconesand molartoothsize[ #'" ]);thenitrogenisotoperecorddemonstratingafully-herbivorouscavebearsintheLatePleistocene [ #'# ];similardamagepatternsle.oncavebearskulls,jaws andlongbonesmainly,inadditiontopunctureandmarks andchewingpresentoncavebearbonesineverycavebear denofEurope[ %# ].( epuncturemarkswereespeciallymisinterpretedtohaveresultedfrom"noncarnivorous"cavebear canines,withround-ovalholesincavebearcublongbones alsomisidenti-edas"Neanderthalpseudo-bone,utes"but thoseresultsarecertainlyfromthemolarcrushingteeth ofhyenas[ %# ].Hyenaswereunabletocrushcavebearcub femoraeandlongbonesintopieces,becausethosewerenot calci-edenough,andonlysubadulttoadultlongboneswere crushedsuccessfullytopiecesinlargeramountsinsome caves[ %# ].( erstevidenceofhyenaimpactsoncavebear populationswaspublishedin!""'basedonbonetaphonomy studiesandfullanalysesofoverlappinghyenaandcavebear densofthePerickCaves[ #'! ].Stilla.erthis,cavebear researchersbelievedin"boneconsumption"ofcavebears, whichwasagainincorrectlypublishedfortheOaseCave(Ro) [ #'& ],wherethehyenaskull[ #* ] -ndwas-rstoverlooked (alsooverlappinghyena/cavebearden).Hyenapresence explainalsotherethelargeamountsofbonefragments(up to!#%ofcavebearbonesdamaged).Hyenasspecialized withoutadoubtinborealmountainforestregionsallover Europe,wherecavebearpopulationswerewellrepresented inpossiblyhigherdensitiesasbelieved("onecavebearfamily percave"[ #'" ]).Evidencefortheircarcassfeeding(possibly alsokillingofcubs=overlapwithlionprey; Figure/ (c)),asa resultofabsenceorrarenessofthemammothanditssteppe megafaunaabundancevariability(especiallyhorse/steppe bison)inthoselandscapes,isdemonstratedforseveralcaves suchasbestillustratedforthePerickCaves[ #'! ]orZoolithen Cave[ $' ]. &$.NeanderthalHumanExhumersandScavengers. ( ere aretwoobviousexampleswhereLatePleistocenespotted hyenasmusthaveexhumedandeatenNeanderthalhuman carcasses.( erstconvincingrecordisfromtheRochelot Cave(F),wheresometeethandcrackedlongbonefragmentswerefoundinaclassicalhyenadenbetweenEemian interglacialequiid/bovidpreyremains[ &$ ].( einterpretationofthenonhumancamporburialinsmallercaves documentstheconsumingofhumansbyLatePleistocene spottedhyenas[ #'$ ].Fracturedboneremainsofseveral NeanderthalhumansfoundintheDivjeBabeICave(Hr) interpretedtohaveresultedfromhumancannibalism[ #'' ] arehereinreconsideredforseveralreasons.Firstly,thecave wasobviouslynotawellstudiedhyenaden(ownobservations oncarnivorebonematerial).Secondly,thehumanlongbone fragmentswhichhavenoevidenceof"hitmarks"suggesting theyresultfromhyenabonecrushingactivities.(emost importantargumentcomesfromcomparingmodernhuman bonematerial,whichwasexhumedfrommorethanone metersindepthbesideahospitalinAfricabyspottedhyenas (unpublishedSutcli+e-modernhyenadencollectioninthe BMNHL),wherethereisevidenceforhumanbodyparts andcraniaimportedintoasmallcaveden.(ere,skulls werecrushedinfragmentssimilarlyasallthelongbones, andthismaterialisakeyfortheunderstandingofhyenas scavengingonNeanderthalsandhumans/apesingeneral. Also,theCroatianDivjeBabeICaveisanotheroverlapping hyenaden/Neanderthalcampsite.Itisplausiblethathyenas exhumedhumanswhichwerenotdeeplyburiedandle. behindonlycraniapieces,teeth,andlongbonefragments. ( ebreakagepatternsandselectivepresenceofcraniaand longbonefragmentscorrelatewiththehyenadensiteof RochelotCave(F)andthemodernAfricandensites,and thereforeNeanderthalsmusthavebeenonthelistofthe hyenaprey,atminimumascarcasses.However,attackson humansbythesepredators,especiallyonlargeportalcaves wherehyenasandhumansoccupiedbothofthose,mustbe expectedbecausebothusedthoseseasonallyatthesameor evendi+erenttimes,whichcomplexsituationhasalsobeen discussedfortheoverlappinghyena/NeanderthalsiteatBalve Cave(D)[ $$ ]. 4.LatePleistoceneSteppeLions (elionsoftheEurasianLatePleistocene(Figures $ % ) werecalledhistorically"cavelions"becausetheywere-rst foundintheZoolithenCave(D)[ ]andothercavesin Europe.However,theywererecentlyrevisedaccordingto DNA[ #'% #'* ]studiestotheLatePleistocenesteppelion Pantheraleospelaea (Goldfuss,#)#").(eholotypeskullhas composedlowerjawsfromotherindividuals[ $% ].( oseLate PleistocenesteppelionsfromtheEemianinterglacialwere smaller(e.g.,modernmaleswerenearlysimilarinsizeto Eemianmales,whichrepresentpossiblyevenanotherwarm periodsubspecies),butthoseoftheWeichselianglacialwere largerusingthelargestskulls[ $% */ ].Similartothemodern Africanlions[ ** ]thecoldperiodmalesintheextinctspecies [ ** ]arefewlarger[ $% ** ],whichisalsore, ectedintheskull recordofEurope,wherassiblingtoearlyadultskullscannot beattributedwelltothesex( Figure' ). ( & .PalaeopopulationsoverEurope. Lionswerethoughttobe "rare"intheLatePleistocenefossilrecordofEurope,but thiswasonlybecauseofalackofresearch,mainly.Inthe past-veyearstheadditionofnewunpublishedmaterialand revisionoflionremainsinGermany[ $% %$ %% %* )& )% ] andCzechRepublic[ %! )! ](#.&*&bones,includingremains of/skeletons)andnew-nds($skeletons)inRomania[ %# %' ] demonstrate"denspalaeopopulations"intheLatePleistocene ( Figure& Table# )withabonerecordratioof#lion/&hyenas. Someindividualskeletons( Figure& )andhundredsofbones havebeendescribedoverthepastyearsfromGermanyand CzechRepublic,demonstratingmorehyenasinthebone record,whichdoesnotre,ecttherealindividualanimal amountsbecausethemortalityandtaphonomysituationfor bothisdi+erent,especiallyinthecavedens.Lionswerefound

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!% PaleontologyJournal insteadlessatopenairsites[ %! %% %* )' ]andaremore abundantincavebearorhyenadens[ $% %# %' )! )$ )% ] ( Table# ). (.#.HolotypeSkullsandSkeletons. ( erecentlyrediscovered holotypeskullof" Felisspelaea "wasoriginallyfoundinthe ZoolithenCaveofBavaria(D)[ ].( elowerjawsfromthis skullareactuallycomposedfromdi+erentindividuals,with onemandiblefromafemaleincorrectlyattributedtothemale skull[ ) ].(elargestpalaeopopulation(possiblyincluding di +erentsubspecies)wasfoundintheEuropeanZoolithen Cave[ $% ].AlsofromthisZoolithenCave,similartoasmany localitiesinEurope,articulatedskeletonsareabsentbecause ofthreereasons:(a)historicallynonprofessionalexcavation andcollecting,(b)rarelioncannibalism(exceptofcubs),and (c)transportionby, oods(thirdposition).Mostrecently,a fairlycompleteskeletonofanindividualsubadultlionesswith braincasedamagewasdescribedfromtheWeichselianglacial SrbskoChlum-Kom nCavehyenaden(CZ)[ )! ]( Figure$ ). Anotherskeletonwithamoreheavilymoderndamaged pelvicarea( Figure$ )ofanotherstrongilllionesswasfound withintheNeumark-NordLake#openairelephantgraveyard sitedatingtotheEemianinterglacial(D)[ %# ].Remains ofthreeglacialagedskeletonsfromtheUrs ilorCavecave bearden(Ro)( Figure$ )[ %' )$ ]arethemostspectacular -ndsatall.Otherglaciallionskeletonshavebeenfoundat variousGerman(Siegsdorf[ ** ],Huttenheim,Edingen,Br uhl [ %% ]),andCzech(Praha-Podbaba[ %& ])openairriverterrace sites.Furtherskeletonshavealsobeenreportedinunclear taphonomiccontextfromSpain(Arrikrutz)[ *$ ]andAustria (Salzburg)[ *' ].(eonlyknowncubskeletonwasfound withinahyenaden(WilhelmsCave,D)andseemtohave beenimportedas"hyenaprey"orwaskilledbyhyenasattheir densiteduringpreystealingorcubkillingpurposes[ )' ]. ( .OntogenyandSexualDimorphism. ( eontogenyin P. l.spelaea wasoriginallyconfusingandyoungindividual materialfromtheZoolithenCavewasincorrectlyattributed toa"cavetiger,"whichwasrevised[ $% ].Cubindividual remainsarerareincontrasttohyenasandhaveonlybeen foundwithapartialskeletonintheWilhelmsCave(D)hyena den[ )' ],whichdemonstratesthatthisanimalwasnota cave"inhabitant",norperiodically"caveuser,"butonlya dweller( Figure) ).Skeletonsareonlyofsubadulttosenilein age( Figure$ ).( esexualdimorphisminmodern[ ** #') #%" ]andPleistocenelions[ $% ** ]iswellknownbestfrom theteeth[ #%" ],crania,butalsoinpostcranialcomparisons [ %% ].Incontrasttohyenas,malelionsareknowntobe largerbasedondatapresentedforseveralcavesandopen airsitesofGermanyandCzechRepublic[ $% ].( ere,the sexidenti-cationremainso.enunclearincubs/subadults, especiallyifthesiteisnotwell-datedtobeinterglacial (smallerforms)orglacial(largerforms)orintheclose overlapofsmallfemalesandlargermales[ $% %% ** ]. ( ( .SteppeLionsasCaveDwellers. Additionaltothetaphonomicandnitrogenisotoperecord,thefossilrecorddataon lionremainsfromcavebeardensindicatethatonlysubadult toadultlionindividualsdiedinthecaves[ $% )$ ].Clearly theuseofcavesbylionscanbeexcludedincomparisonto themortalityofcavebearsandhyenas( Figure) )[ % ].Late Pleistocenesteppelionswereopenenvironmentinhabitants rangingintheirterritoriesfrommammothsteppetoboreal forestenvironments,similarinethologytotheirmodern relativesinAfrica[ $% %% )$ ].(erefore,thehistorically named"cavelion"wasrecentlyrenamedthe"steppelion"[ ) ] accordingtoDNA,taphonomic,andbehaviouralinformation. (.%.ReindeerandBisonHunters. Nitrogenisotopicstudies ontheLatePleistocenesteppelionsindicatethattheyseem tohaveswitchedtoreindeerhuntingattheendoftheLate Pleistocene[ #%# ]asaresultofextinctionofthemammoth steppemegafaunaandcavebears[ #') ].(isiscomparableto modernlionswhichtypicallyhuntgazelles[ #') #'/ ].Lions, asfastrunners,seemtohavespecializedmoreinfastprey suchascervids( Cervus,Megaloceros, and Dama )orbovids ( Bos Bison )intheLatePleistocenesimilartothelargewolf subspecies,possiblybothhuntinginpacks.Itmustalsobe expectedthattheraresteppebison/aurochsremainsinmost hyenadensre,ectthepreycompetitionwithlions/wolves.As knowninAfrica,bovidsarethemaintargetoflionsinmany regions[ #') ]. (.".MammothandForestElephantInitialFeeders. Specializedpredationonyoung-subadultelephants[ #%! ]cannot bedemonstratedintheLatePleistocenerecordofhyena denboneassemblages.Atthesesitesthemammothpresence/absencedependsmoreonthelandscapemorphology [ '* ]( Figure/ (b)).Informationofthelion'slargestpreycomes bestfromthefossilelephantcarcasssitesSiegsdorf( M. primigenius skeleton,glacial)andtheNeumark-NordLake # site( Palaeoloxodon skeletons,interglacial).Asdocumented inAfrica,modernlionsstarttoconsumethetrunk,andfrom theanusfromwheretheyeattheintestinesofthebodycavity nallyfrominside[ ') ].( isisalsowellshownfrommodern hyenasandlionsindocumentary-lms[ #$) ]andamonitored Loxodonta carcass[ #$* ].Modernhyenasfeedonelephant carcassesatthekillingsiteinthecarcassdecompositionstage # [ #$* ]. ( $.CaveBearCubKillers. IncontrasttotheLatePleistocene spottedhyenas,thepreyofthelionsismuchmoredi:cultto reconstruct,becausetheydidnot[ #') #'/ ]accumulateprey orboneremainsoftheirpreyordonotleaveconsistenlydamagedbones.(etheoryaboutlionskillingcavebearseven indeepcavesresultedfromtaphonomicstudies,whereas convincingevidencehasrecentlyemergedfromUrs ilorCave cavebearden(Ro, Figure$ )[ %# %' ].Furtherevidencefrom taphonomicstudiesisdemonstratedfromseveralcavesin theSauerlandKarst[ )% ].Similarresultswerefoundnally attheZoolithenCavewiththelargeststeppelionpopulation foundeverinalargecavebearden(aboutahalfmillion cavebearbones),whereafeedingspecializationoncavebears wasarguedduetotheabsenceofmammothsteppefaunain mountainousregions[ %# ].(elatestnitrogenisotopedata

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PaleontologyJournal !* supportsthecavebearhuntinginterpretation,whereasothers evenwentfurthertodeclarethatonly"cubs"wereamain foodsource[ #%# ].MorerecenttaphonomicstudiesatSophie's Cavesupportthis.(ehuntingofcavebearsbylionsexplains thatcavebearshibernatedasdeepaspossibleinlargercave systemsevenindangerouspassagestoprotectthemselves againstlionattacks[ $% ].Huntingcavebearsinthecaves seemstohavebeendonemainlybyfemales.(erearesome largercavebeardensinEurope(e.g.,KepplerCave,Urs ilor Cave,andpossiblyalsoHermann'sCave),wheretwothirdof thelionbonematerialbelongstofemales.(isisasimilar behaviourasinmodernlions,wheredominantlionesseshunt inpacks[ #') #'/ ].ApairorpackhuntbyLatePleistocene steppelionsinthecavesmustbeexpected. (.) .DeathlyBattleswithCaveBears. ( elionbonerecord incavesmightsimplydemonstratethatfemalesweremore easykilledbygrownupcavebearsduringcon,ictsinthecave dens[ %' ].Interestinglytheamountatoverlapathyena/cave beardensitesofthemale/femaleratioisoppositetothatof lions(BilsteinCave[ )& ],PerickCaves[ %$ ],andZoolithen Cave[ $% ])withonlyonethirdofthebonesfromfemale hyenas.Lionsmusthavebeenunsuccessfulsometimeswhile huntinginthecave( Figure) (d)).( erearethreeexamples oflionskeletonswhichseemtobetheresultoflost-ghts: (a)SloupCave"skeletons"(atminimum! )[ %! ],(b)Urs ilor Caveskeletons(atminimum&, Figure% ),and(c)Zoolithen Cave[ $% ](unclearamount,butremainsarefromoriginally partiallyarticulatedskeletons[ $% ]).(elionremainsin bone-richcavebeardensneverexceed#&%o+thetotal NISPboneamount[ $% %# %* )& )$ ].( iscalculatestoonly onedeadlionper#", """yearsatmaximum,whichsuggests o .ensuccessfulhuntingonthe"hibernating"andeasyto killcubsovertenthousandsofyears.(eaforementioned "normalmortality"incavebearsofEurope[ #%& #%' ]does notexist;instead,thehighmortalityincavebearcubs(also hereZoolithenCave: Figure) (a))seemstobeacombination ofboth:naturaldyingandpredationmortality( Figure) ), whichisdemonstratedbythepercentageofbitedamaged cubbones,whereasmostofthebitedamagesclearlyareof postmortaltimes(e.g.,chewedjoints).Alsothepictureofthe cavebearslifeanddeathwaspresentedincorrectly[ #'" ],such asinthebonetaphonomy(cavebearsas"cannibals"[ #%% ]), bynottakingthepredatorsintoanyaccountinanycavebear populationstatistics/mortalityanalyses.However,becauseof thelackofmodernlion/bearpopulationoverlap,thereare nopossibilitiestocomparethisveryuniquesituationoflions huntingcavebearsdeepincavesduringtheLatePleistocene. (.!.PermanentWarwithHyenas. Modernspottedhyenas (theleadingfemale)arenormallykilledbytheleadinglion whilemalelionsarerarelykilledbyhyenas[ #$) ].Late Pleistocenespottedhyenaswerefeedingatleastondeadlions (assimilarastheirmodernrelatives)andimportedtheir remainstocaveandopenairdensites(alsosimilartomodern [ /$ ])whichiswelldocumentedintheLatePleistoceneof Germany[ %$ %* ]( Table# ).( econ, ictsare/wereabout territory,cubprotection,andprey[ /' #') ],butintheLate Pleistocene,battlesinhyenadencavesaremorefrequent, whichhasonlylimitedmoderncomparisonduetolackof extensivecavespresentinAfrica.(ecave"battlemodel"of theLatePleistocenepredatorsestablishedfortheZoolithen Cave[ $' $% ]( Figure) (b))demonstratesthecaveuseby hyenas(entranceareaasden)andcavebears(deeperfor hibernation).Itremainsunclear,ifbothsynchronouslyused cavebranchesatleastatsomelargecavesystems[ $' ].Inthe ZoolithenCave,skullbitetraumapathologiesarepresentat allthreeanimals,hyenas,lions,andcavebears( Figure) (b)) [ %) ].Alsoamisinterpretationwasmadebeforehere[ #%* ]that thedamagetocavebearskullsresultedfrom"Neanderthal hunts"[ #%) ]whichsuggestednottohaveoccurredatall inEuropefortheMiddlePalaeolithic,butonlylaterwith newweapontechnologyofmodernhumansintheAurignacian/Gravettian(projectilefragmentinvertebra,cutmarks oncavebearbones,projectilesdeepincavebeardens,and pathologicalcavebearboneswithboneprojectiledamage) [ #%/ #*" ].(ecranialdamageontheZoolithenCavelion skullissimilartothatwhichhasalsobeenfoundonother skullsfromcavebeardens(SloupCave)andhyenadens(SrbskoChlum-Kom nCave)inCzechRepublic( Figure* )which arealsoattributedtobattlesbetweenlionsandcavebearsor betweenlionsandhyenas[ %! %) ].Inalltheseoverlapping hyena/cavebeardensthereseemstohavebeentwomain "warzones."(e-rst(zone#)isattheentranceofahyena den(battlewithhyenaswhilestealingprey,orkillingtheir cubs),whichresultedalsoinlionkillsbyhyenas,whichisbest demonstratedbytheillyounglionessskeleton-ndfromthe SrbskoChlum-Kom nCavehyenapreydepotden( Figure$ ) [ %! ].Itwasproposedthattheyounglionesswithabitetrauma damage,whichwaspossiblyexcludedfromthepride,tried tostealpreyfromahyenadenpreystoragesiteandwas killedbythehyenaclan(orithasbeenimportedascomplete carcass)[ %! ].Asimilarbutdi+erentsitetype(scavenging openairsite)battlesituationwaspresentedfortheilllioness fromNeumark-NordLakeI,whichhadonlyonelostupper jawcanineandhindlegtraumaillness.(ere,ascenario isreconstructedwheretheoutnumberedweaklionesswas killedbecauseofpreybattleswithahyenaclan[ %! ].Most ofthecranialtraumadamageseemstohaveresultedby thesebattlesandnotfromcavebearbattles,whichalsomust havehappened.(osecanbedeterminedbythepathologicalbonesofbothtoppredators( Figure* ).Newlystudied traumaticpathologiesinlions[ %) ]andhyenas(herein)are di +erent( Figure* )andareevendi+erentfromthoseofthe cavebears,whichlatterabundantandverydi+erenttypes ofpathologiesmustberevisedandstudiedinfuturewith predationandhuntingbackground.Asalreadydiscussedfor lions[ %) ],bitedamagecausingexostosesbonegrowthare causedbylionsandarefoundmainlyontheirskullsaggital crest,thefore,andmuchlessthehindlimbs( Figure* )[ %) ]. Lionsstrugglewiththepreywiththeforelimbsandattack mainlyontheheadofthecarnivoreantagonist[ %) #') ]. ( isexplainsthedamageonhyenaskullstoo,includingon theirsaggitalcrestashasbeenfoundonatleastsixskulls ( Figure* ).Asmentionedintheskullshapetypesbefore, healinga.erstrongbitedamagescausesstrong-convexcrests suchasseenatremainsfromtheZoolithenCaveand

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!) PaleontologyJournal Br uhl-Spieswiesenopenairsites.InLatePleistocenespotted hyenas,othertraumapathologiesontheshoulderareaare alsofoundinasingleexample(SrbskoChlum-Knom n Cavealsopossiblebattlewiththeaforementionedlioness) andmosto.enontheirdistalhindlegs( Figure* ).In comparison,modernAfricanspottedhyenasinsteadattack lionsandotherhyenasbybitingthemonthehindlegs. (efactthattheyaremuchmorecommononhyenasthan lionhindlimbsdemonstratesthattheseareresultsfrom mainlyintraspeciescon, ictsofhyenas.(ewarbetween lionsandhyenasisalsodocumentedwithremainsfound inlargerhyenadenboneassemblages(see Table# )where bonesmightresulteitherfromakilloronlyscavengedcarcass whichhyenasfoundandimportedtotheden.Comparedto modernAfricanlionandhyenas,theirbattlesresultin'"% ineachspeciesmortality[ #$) #*# ],whichmustbeexpected tobesimilarbutwithcavebearmortalityimpactintheLate Pleistocene. ( &*.LatePleistoceneSteppeLionCannibalism. IntheLate Pleistocene,therewasalsothewar(zone!deepinthe cave)withthecavebears.(eseattacksweremademainly oncubs( Figure* )andfromtimetotimelionsmusthave beenkilledbythemaleorfemalesprotectingtheircubs. ( esecarcasseswerenotconsumedbyherbivorouscavebears whichexplainswhylionskeletonsincavesareo.encomplete, suchasthosefoundinUrs ilorCave( Figure% ).However,one ofthesesubadultlionessskeletonswasscavengedbyalarge toppredator(hyenaorlion)assuggestedbythebitemark sizes.( eslightdamageonbonejointsonlyandabsenceof crackedbonesmightindicateacannibalisticscavengingbya lion,ratherthanbyahyena.Inthiscaveareaabout)""meters farfromtheentranceinasecondcavelevelwherelionswere possiblytrapped,andpossiblythissituationinUrs ilor,Cave istheonlyindicationforpossiblelioncannibalismunder stresssituations,whichstresscannibalismwasalsorarely observedinmodernAfricanlions[ #') ].Ifthisistrue,thenwe haveanotherargumentfor"lionshuntinginpacksorpairs" whereastheyoungestandweakestonewas-nallythetarget ofotherlions. 5.Conclusions LatePleistocenespottedhyena Crocutacrocutaspelaea (Goldfuss,#)!&)andsteppelion Pantheraleospelaea (Goldfuss, #)#")arerepresentedinCentralEuropebyaratioof & hyenas/#lionremainwhichisestimatedonmaterialfrom #"%mainlycaveandinlesseramountofopenairsites.Oneto threepercentofthelionremainsathyenadensitesindicate atminimumtheirscavengingbyhyenas,whichpossiblyeven killedtimebytimecubs,adolescents,lioneses,or/andweak individuals,similaraswellreportedbytheAfricanrelatives. ( eextantlasthyenasandlionsofEuropehavesimilarities intheirecologiestomodernAfricanhyenas/lionsresulting incompetitionaboutpreyandterritory,whereasonlyhyenas usedcavesasdensites.Howevertherearedi+erencesto modernAfricanhyenas/lions,becausetherecavesareless abundant,andcavebears(orotherbears)areabsent,asare thepreygroupofcervids.CannibalismwithinLatePleistocenespottedhyenasisdocumentedatmanydens,whereas onlytwoindividualskeletonsfromEuropere, ectspecial taphonomicsiteconditions(vertical-diagonalsha.sorprey storagesites).LatePleistocenespottedhyenasle.bone accumulationsatthreedi+erentdentypes:(# )birth/natal dens,(!)communaldens,and(&)preystoragedens.Noneof thelionandwolfcompetitorsproducedboneaccumulations, norsuchmassivebonedamagesfoundwithinhyenaden boneassemblages.Onlyhyenasdevelopedatallmagamammalgroupsasimilarbutcheringtechnique,todecompose carcasses,whosebodyparts(evenNeanderthal/Cromagnon humanremainsofexhumedshallowburials)wereimported todenstoavoidcon,ictswithotherpredators.Feeding specializationsbyhyenas,lions,andwolvesondi+erent megamammalgroupspartlyoverlap(e.g.,cavebears).Lions andwolveswerespecializedcervid(fastrunningprey)and bovid(largeherds)hunters,whereashyenaswerethemain meatcolossusscavengersandcarcassdecomposersofwoolly mammothandwoollyrhinoceros,whichsubadults/adults haveevenforhyenasuncrushablebones.LatePleistocene spottedhyenasle.repeatedlysimilardamagedlargeprey bonesalloverEurope.Similartomodernspottedhyenas,Late Pleistoceneonesareexpectedtohavehuntedinclansequids (PrzewalskihorsesintheLatePleistocene-zebrasmodern). FoetalPrzewalskihorseremainsprovequiteuniquelythe huntinspringtimesatoneCzechhyenacavepreystorage andcommunaldensite.Steppelionsasopenenvironment (mammothsteppetoborealforests)felidsfocusedoncervids withreindeertargetingspecializationduringtheLastGlacial Maximumaround!!."""BP.,whenothermegafaunabecame rarerorextinct.(eyalsomusthavehuntedpossiblyin packsbovids(steppebison/aurochs)orsaigaantelopesinthe steppeorvalleyenvironments.LatePleistocenelionsnever usedcavesevenforshort-termperiods;theyonlydwelled forcavebears,mainlycubhunting.Allthreetoppredators (lions,hyenas,andwolves)fed(huntedorscavenged)oncave bearsinborealforestofmiddlehighelevated(#'"%'"a.s.l) mountainregions(lionsevenuptoelevationsof# '""a.s.l.) whichcauseddeathlycon,ictsincavesbetweenallofthem (inter-/intraspeciesghts)thathavenomodernanalogue. ( osebattlescausedbitedamagesespeciallyontheirskulls andlegs,whichproduceddi+erentpostcranial,butsimilar cranialbitedamagepathologies.Someofthebitemarksalso foundatlionskeletonsandbonescannotbeseparatedclearly intheiraliveorpostmortalorigin. ConflictofInterests (eauthordeclaresthatthereisnocon, ictofinterests regardingthepublicationofthispaper. Acknowledgments ( eauthorthanksallthemuseumsandinstitutionsmentionedandtheircuratorsfortheirsupporttostudybone materialfromvarioussitesinGermany,CzechRepublic, Austria,andEngland.(ecompanyandprivateresearch

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PaleontologyJournal !/ institutePaleologic( www.paleologic.eu )sponsoredthesciencetothe"EuropeanIceAgeSpottedHyenaProject."Arst criticalreviewandspellcheckwasmadethankfullybyDr.G. ZazulafromtheBeringeriaCenter,Canada.Dr.M.Sabolalso commentedona-rstdra..J.McNamara(WorldMuseum ofMan,www.worldmuseumofman.org)supportedthepaper withphotosoftheAustrianBadelCavelionandhyena skullsandmadeapaperspellcheck.FinallyG.Teichmann contributedthehyena,lion,andcavebearillustrations. References [ # ]J.F.Esper,"Ausf uhrlicheNachrichtvonneuentdeckten Zoolithenunbekanntervierf u§iger(iere,unddenensie enthaltenden,sowieverschiedenenanderndenkw urdigen Gr u .enderObergeb urgischenLandedesMarggra.hums Bayreuth,"Knorrs,N urnberg,Germany,#**$. [ ]C.G.Diedrich,"(erediscoveredcavebear" Ursusspelaeus Rosenm uller#*/$"holotypeoftheZoolithenCave(Germany) fromthehistoricRosenm ullercollection," ActaCarsologica Slovacia ,vol.$*,no.#,pp.!'&!,!""/. [ & ]J.C.Rosenm uller,"Quedamdeossibusfossilibusanimalis cuiusdam,historiameiusetcognitionemaccuratioremillustrantia,dissertatio,quamd.!!.Octob.#*/$addisputandum proposuitIoannesChrist.Rosenm ullerHe§berga-Francus, LL.AA.M,"in +eatroAnatomicoLipsiensiProsectorAssumto SocioIo.Chr.Aug.HeinrothLips.Med.Stud ,pp.#&$,Cum tabulaaenea,Leipzig,Germany,#*/$. [$]G.L.C.F.D.Cuvier,"SurlesossementsfossilesdesHy ` enes," AnnalesduMus eeHistoireNaturelleParis ,vol.%,p.#!*, #)"'. [']G.A.Goldfuss,"DieUmgebungenvonMuggendorf.Ein Taschenbuchf urFreundederNaturundAltertumskunde," Erlangen,Germany,#)#". [%]G.A.Goldfuss,"OsteologischeBeitraegezurKenntnisverschiedenerSaeugethierederVorweltIV.UeberdenSchaedel desHoehlenloewen," VerhandlungenderKaiserlichenLeopoldinischenCarolinaeischenAkademiederNaturfreunde ,vol.#", no.!,pp.$)/$/$,#)!#. 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